Skip to main content

Full text of "SIDA, contributions to botany."

See other formats 


S1DA CONTRIBUTIONS TO 
BoTANy WAS FOUNDED BY 
Lioyo H. SHINNERS (LEFT) 
IN 1962. INHERITED BY 
WILLIAM F. MAHLER 
(RIGHT), DIRECTOR EMERITUS 
oF BRIT in 1971, AnD 
SINCE 1993, IT HAS BEEN 


PUBLISHED BY Brit Press. 


Barney L. Lipscoms, EDIToRr 

Botanical Research Institute of Texas 
509 Pecan Street 

Fort Worth, Texas 76102-4060, US, 
817 332-444] / 81 


’ 332-4112 FAX 


Electronic mail: sida@brit.org 


Home page at the URL: http://www.brit.org/sida/ 


W. 


{JOHN W. THIERET, ASSOCIATE EDITOR 
Dept. of Biological Sciences 
Northern Kentucky University 


Highland Heights, KY 41099 USA 


FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR 


Dpto. de Botanica, Facultad de Biologia 


Universidad de Lean 


E-2471 Leon, SPAIN 


The views ex 


reflect those of the editors or of the Botanical 
Research Institute of Texas 


Guidelines for contributors are available upon request 


and on our SIDA home page as well as the last pages 


of each volume. 


Subscriptions for year 2006: 
$39. Individual 
$75. USA Institutions 


$85. Outside USA 


numbers issued twice a yeal 


< 
N 
zx 


BOTANICAL RESEARCH INSTITUTE OF 


ISSN 0036-1488 


yressed in this journal do not necessarily 


TABLE OF CONTENTS 

DEDICATION 

John W. Thieret (1926-2005) 

RALPH L. THOMPSON 

John W. Thieret, the curator 

Rosert F.C. NAczi 

John W. Thieret, a student's perspective (1967-2005) 
Rosert R. HAYNES 

John W. Thieret, valuable botanical friend (1965-2005) 
RONALD L. STUCKEY 

John W. Thieret, colleague and editor friend (1969-2005) 
WILLIAM F. MAHLER 

John W. Thieret, associate editor of Sida (1972-2005) 
BARNEY LipscOMB 

SYSTEMATICS 

A new narrowly endemic species of Clematis (Ranunculaceae: sub- 
genus Viorna) from northeastern Texas 

Dwayne ESTES 


Hill cane (Arundinaria appalachiana), a new species of bamboo 
(Poaceae: Bambusoideae) from the southern Appalachian Mountains 
J.K. TRripcett, A.S. WEAKLEY, AND L.G. CLARK 

Revision of Bauhinia subgenus Bauhinia section Amaria (Cercideae: 
Caesalpinioideae: Fabaceae) 

RICHARD P. WUNDERLIN 

Dendrophorbium restingae (Asteraceae: Senecioneae). anew species 
from Sao Paulo, Brazi 

AristOnio M. TELEs, Jimi N. NAKAJIMA, AND JOAO R. STEHMANN 


Novedades en Gentianaceae para América del Sur 
Eva M. FicippA AND GLORIA E. BARBOZA 


Amelichloa: a new genus in the Stipeae (Poaceae) 
Mirta O. ARRIAGA AND Mary E. BARKWORTH 


Taxonomy of the Cuscuta pentagona com plex (Convolvulaceae) 
in North America 
Minal Costed, Guy L. NESOM, AND SASA STEFANOVIC 


Taxonomy of the Cuscuta salina-californicacom plex (Convolvulaceae) 
Minal Costea, Guy L. NESoM, AND SASA STEFANOVIC 
Taxonomy of C tag d Cuscuta umbrosa(C olvulaceae) 
Minal Costa, Guy L. NESoM, AND SASA STEFANOVIG 


Taxonomy of the Cuseuta indecora (Convolvulaceae) complex in 


North America 
Minal Costea, Guy L. NEsoM, AND SASA STEFANOVIC 


21 


25 


33 


55 


65 


97 


123 


129 


145 


A new species of Pediomelum (Fabaceae) from the lower Piedmont 
Plateau of Georgia and South Carolina 

JAMES R. ALLISON, MICHAEL WAYNE Morris, AND ASHLEY N. EGAN 
Big-fruited buckthorn, Sideroxylon macrocarpum (Sapotaceae), a 
long-forgotten Georgia endemic 

JAMES R. ALLISON 

Sexual and apomictic prairie {leabane (Erigeron strigosus) in Texas: 
geographic analysis and a new combination (Erigeron strigosus 
var. traversii, Asteraceae) 

RicHARD D. Noyes, HEIDI GERLING, AND CARLA VANDERVOORT 

More American black sapotes: new Diospyros (Ebenaceae) for 
Mexico and Central America 

MITCHELL C. PROVANCE AND ANDREW C, SANDERS 

Taxonomy of North American species of Oldenlandia (Rubiaceae) 
Epwarb E. TERRELL AND HAROLD ROBINSON 


New taxa and combinations in cultivated bamboos (Poaceae: 
Bam busoideae 

C.M.A. STAPLETON 

Estudios en las Apocynaceae neotropicales XXIV: ees eee 
deae: Echiteae), un desapercibido nuevo género endémico de Bahia, 
Brasil 

J. FRANCISCO MORALES 


Estudios en las Apocynaceae neotropicales XX V: novedades y nuevos 
reportes en las Apocynaceae (Apocynoideae: Rauvolfioideae) de 
Venezuela 

J. FRANCISCO MORALES 


Taxonomic overview of the Heterotheca villosa complex (Asteraceae: 
Astereae) 

Guy L. Nesom 

Taxonomy of Lantana sect. Lantana (Verbenaceae): |. correct 
application of Lantana camara and associated names 

ROGER W. SANDERS 

Crataegus series Parvifoliae and its putative hybrids in the south- 
eastern United States 

J.B. PHipps AND K. Dvorsky 

A new species of Stenanthium (Melanthiaceae) from Tennessee, U.S.A. 
B. EUGENE WOFFORD 

A new species of Dodecatheon (Primulaceae) from the northern 
coast range of Oregon and Washington 

KENTON L. CHAMBERS 


The lectotypification and 19" century history of Croton alabamensis 


(Euphorl biaceae s.s. ) 


KENNETH J. WURDACK 


Two new species of Elymus (Poaceae) in the southern U.S.A. and 
other notes on North American Elymus species 

JULIAN J.N. CAMPBELL 

Anew hybrid genusand 12 new combinationsin North American grasses 
Mary E. BARKWORTH 

Quadruple, triple, double, and simple pappi in the goldenasters, 
subtribe Chrysopsidinae (Asteraceae: Astereae) 

JOHN C. SEMPLE 

Notes on types in Apocynaceae: Asclepiadoideae in Cuban herbaria 
and four lectotypifications in West Indian Gonolobinae 
ALEXANDER KRINGS AND PAUL R. FANTZ 

Comments on the Gerbera-complex (Asteraceae: Mutisieae) 

Hans V. HANSEN 

Validation of the name Dahlia sublignosa (Asteraceae) 

DayLe E. SAAR AND PAuL D. SORENSEN 

A new combination in Polytrichastrum (Bryophyta: Polytrichaceae) 
G.L. SMITH MERRILL 

A new combination in Tetraphis (Bryophyta: Tetraphidaceae) 
JupitH A. HARPEL 


A new combination in the Bromus catharticus complex (Poaceae: 
Bromeae sect. Ceratochloa) 

ANA MariA PLANCHUELO 

CHROMOSOME NUMBERS 

Chromosome numbers for western and arctic North American 
species of Antennaria (Asteraceae: Gnaphalieae) 

JERRY G. CHMIELEWSKI 

ANATOMY AND MORPHOLOGY 

Root-shoot anatomy and post-harvest vegetative clonal develop- 
ment in Lophophora williamsii (Cactaceae: Cacteae): implications 
for conservation 

MArtTIN TERRY AND JAMES D. MAUSETH 

ETHNOBOTANY 

Taxonomy and conservation of medicinal plants in canal-irrigated 
areas of Punjab, Pakistan 

KHALID FAROOQ AKBAR AND MOHAMMAD ATHAR 

BOTANICAL HISTORY 

Notes on the life and work of James Brigham McFarlin, Florida botanist 
TOM PALMER 


FLORISTICS, ECOLOGY, AND CONSERVATION 

Floristic diversity and composition of terra firme and seasonally 
inundated palm swamp forests in the Palma Real Watershed in 
lower Madre de Dios, Peru 


FERNANDO H. CorRNEJO VALVERDE, JOHN P. JANOVEC, AND MATHIAS W. TOBLER 


A vascular plant inventory of Starkey Wilderness Preserve, Pasco 
County, Florida 

EMILY FERGUSON AND RICHARD P WUNDERLIN 

Vascular floras of Sonoita Creek State Natural Area and San Rafael 
State Park: Arizona’s first natural-area parks 

STEVEN P. MCLAUGHLIN 

Annotated checklist of the vascular flora of the Big Sandy Creek 
Unit, Big Thicket National Preserve, Texas 

Larry E. Brown, BARBARA R. MAcRoserts, MICHAEL H. MACROBERTS, 
Pau. A. HARCOMBE, WARREN W. Pruess, I. SANDRA ELSIk, AND 

STANLEY D. JONES 

The north-south transition of flora across Arkansas: a preliminary 
phytogeographical analysis 

MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS 

High levels of seed inviability among seven populations of the en- 
dangered Short’s goldenrod (Solidago shortii: Asteraceae) 

PATRICK J. CALIE, ROBERT EC. NAcCZ1, CHRISTINA SHACKLEFORD, AND 


JESSICA CAICEDO 


Naturalization and extirpation of water hyacinth (Eichhornia 
crassipes, Pontederiaceae) in southwestern Arkansas, U.S.A. 

RENN TUMLISON AND BRETT SERVISS 

Fungi associated with Borrichia frutescens (Asteraceae): insect galls 


and endophytes 
DIANE TE StRAKE, AMY HADDOCK KEAGY, AND PETER D. STILING 


Type localities of vascular plants first described from Ohio: supplement 
JAMES S. PRINGLE 

Notes on southwestern Moraceae 

ALAN T. WHITTEMORE 

Exotic plant introduction in Kansas, two new species 

IRALEE BARNARD 

New and disjunct records of Eleocharis liesneri (Cyperaceae) from 
South America 

Davip J. ROSEN 

New and significant records of vascular plants for Florida and for 
Collier County and Lee County, Florida 

GEORGE J. WILDER AND MArtHA R. McComprs 

Carex bicknellii (Cyperaceae) new to Arkansas 

PauL M. McKenzie, C. THEO WITSELL, AND JOE WoOoLBRIGHT 


Green fringed orchid (Platanthera lacera, Orchidaceae) in south- 

ern Louisiana 

CHARLES ALLEN, SARA THAMES, HowARD ANDERSON III, Bit NEWTON IIL, 

RHONDA HAMPTON, AND GEORGE FISHER 805 


Seymeria falcata (Scrophulariaceae), a new record for Texas and 

the United States 

JOSELYN FENSTERMACHER 811 
Cayratia japonica (Vitaceae) new to North Carolina and an updated 

key to the genera of Vitaceae in the Carolinas 

ALEXANDER KRINGS AND Rosert J. RICHARDSON 813 
New and noteworthy collections for Arkansas 

JAMES H. PECK AND BreTT E. SERVIss 817 


Clematis morefieldii (Ranunculaceae) new to Tennessee 

Dwayne ESTES AND CHRIS FLEMING 821 
Gratiola brevifolia (Plantaginaceae) new to the flora of Delaware, 

the Delmarva Peninsula, and the Mid-Atlantic 

WESLEY M. KNAPP AND DWAYNE ESTES 825 
Hedyotis australis (Rubiaceae) new to Missouri and Florida and 

related species in the south-central United States 

WALTER H. LEwis 831 
Book reviews and notices 144, 150,176, 196, 208, 226, 242, 330, 354, 366, 422, 
446, 460, 468, 484, 502, 532,538, 544, 546,548, 564, 614, 634, 660, 764, 768, 776, 
780, 800, 810, 816, 830, 837 

Announcements 78, 96, 724 


Index to new names and new combinations in Sida 22(1), 2006 

Achnatherum arnowiae (S.L. Welsh & N.D. Atwood) Barkworth, comb. nov-—496 

Amelichloa Arriaga & Barkworth, gen. nov—146 

Amelichloa ambigua (Speg.) Arriaga & Barkworth, comb. nov—147 

Amelichloa brachychaeta (Godr.) Arriaga & Barkworth, comb. nov—147 

Amelichloa brevipes (E. Desv.) Arriaga & Barkworth, comb. nov—147 

Amelichloa caudata (Trin.) Arriaga @ Barkworth, comb. nov—148 

Amelichloa clandestina (Hack.) Arriaga & Barkworth, comb. nov.—148 

Ammophila breviligulata subsp. champlainensis (F Seym.) Walker, Paris & 
Barrington ex Barkworth, comb. nov—496 

Arundinaria appalachiana Triplett, Weakley & L.G. Clark, sp. nov—88 

Bahiella J.F Morales, gen. nov.—342 

Bahiella blanchetii (A. DC.) J.F Morales, comb. nov—342 

Bahiella infundibuliflora J.F. Morales, sp. nov—345 

Bauhinia amatlana Wunderlin, sp. nov—99 

Bauhinia arborea Wunderlin, sp. nov—102 

Bauhinia ayabacensis Wunderlin, sp. nov—105 

Bauhinia geniculata Wunderlin, sp. nov—111 

Bauhinia petiolata var. caudigera (S.F Blake) Wunderlin, comb. nov—115 


Borinda angustissima (T.P. Yi) Stapleton, comb. nov—332 

Borinda contracta (T-P. Yi) Stapleton, comb. nov—332 

Borinda nujiangensis (TP. Yi) Stapleton, comb. nov—332 

Borinda utilis (TP. Yi) Stapleton, comb. nov-—332 

Bromus catharticus var. elata (E. Desv.) Planchuelo, comb. nov-—556 
Clematis carrizoensis D. Estes, sp. nov—67 

Cuscuta indecora var. attenuata (Waterfall) Costea, comb. & stat. nov—216 
Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, comb. & stat. nov—545 
Dendrophorbium restingae A. Teles, ].N.Nakaj. & Stehmann, sp. nov—123 
Diospyros costaricensis M.C. Provance & A.C. Sanders, sp. nov—291 
Diospyros gomeziorum M.C. Provance @ A.C. Sanders, sp. nov.—282 
Diospyros tuxtlensis M.C. Provance G A.C. Sanders, sp. nov—295 
Dodecatheon austrofrigidum K.L. Chambers, sp. nov—462 


— 


Drepanostachyum falcatum var. sengteeanum Stapleton, var. nov—332 

<Elyleymus hultenii (Melderis ex Hultén) Barkworth, comb. nov-—496 

<Elyleymus mossii (Lepage) Barkworth, comb. nov-—497 

<Elyleymus ontariensis (Lepage) Bark worth, comb. nov.—497 

Elymus X cayouetteorum (Boivin) Barkworth, comb. nov—498 

Elymus churchii J.J.N. Camp., sp. nov—486 

Elymus lanceolatus subsp. riparius (Scribn. & J.G. Smith) Barkworth, comb. & 
Stat nov.—498 

Elymus texensis ].J.N. Camp., sp. nov-—488 

Erigeron strigosus var. traversii (Shinners) Noyes, comb. & stat nov—273 

Fargesia apicirubens Stapleton, sp. nov—331 

Gentianella fabrisii Filippa et Barboza, sp. nov—130 

Heterotheca polothrix Nesom, nom. et stat. nov— 373 

Heterotheca sierrablancensis (Semple) Nesom, comb. & stat. nov—374 

Indocalamus hamadae (Hatus.) Stapleton, comb. nov.—332 

Lantana camara subsp. aculeata (L.) R.W. Sanders, comb. & stat. nov—394 

Lantana nivea subsp. mutabilis (WJ. Hook.) R.W. Sanders, comb. & nov—395 

Lantana strigocamara R.W. Sanders, sp. nov—392 

<Leydeum littorale (HJ. Hodgs. @ W.W. Mitch.) Bark worth, comb. nov—498 

Leymus californicus (Bol. ex Thurber) Bark worth, comb. nov.—498 

Mandevilla angustata (Steyerm.) J. Morales, comb. & stat. nov-—356 

Mandevilla tristis J.) Morales, sp. nov—359 

<Pascoleymus Barkworth, gen. hybr. nov—499 

<Pascoleymus bowdenii (Boivin) Bark worth, comb. nov—499 

Pediomelum piedmontanum J.B. Allison, M.W. Morris & A.N. Egan, sp. nov—229 

Polytrichastrum alpinum var. sylvaticum (Menzies) G.L. Merrill, comb. nov—547 

Pseudoroegneria spicata [. inermis (Scribn. & J.G. Sm.) Barkworth, comb. nov. & 
Stat. nov—499 

Pseudoroegneria spicata {. pubescens (Elmer) Barkworth, comb. nov. & stat. 
nov.—499 

Sideroxylon macrocarpum (Nutt.) J.R. Allison, comb. nov—245 

Stenanthium diffusum Wofford, sp. nov— 

Tetraphis pellucida var. trachypoda (Kindb. ex Paris) Harpel, comb. nov—551 


JOHN W. THIERET, ASSOCIATE EDITOR 
FROM 1972-2005 


This issue of Sida is dedicated to you 
and your enormous contributions to the journal! 


For all that you’ve done | thank you! 
Barney Lipscomb, Editor 


TO EVERYTHING THERE IS A SEASON 
AND A TIME TO EVERY PURPOSE UNDER HEAVEN: 
A TIME TO BE BORN AND A TIME TO DIE; 
A TIME TO PLANT AND A TIME TO PLUCK UP THAT 
WHICH IS PLANTED ... 


Preceding page: John Thieret at his book signing of National Audubon Society 
Field Guide to North American Wildflowers: Eastern Division in 200L 


JOHN W. THIERET (1926-2005) 


Ralph L. Thompson 


Berea College Herbarium 
Biology Department 

Berea College 

tucky 40404-2121, U.S.A. 


Berea, | 


ABSTRACT 


John W. Thieret (1926-2005), an internationally aan American plant taxonomist, is remem- 
dasaconsummate field botanist, exemplary teacher, acclaimed author, superb editor, fine her- 


er 
barium director, inspiring mentor to students ae ee and noble friend. 


RESUMEN 


John W. Thieret (1926-2005), tax6nomo de plantas estadounidense reconocido en todo el mundo, es 


Qu 


recordado como un consumado botanico de campo, un profesor ejemplar, un autor aclamado, un 
magnifico redactor, un excelente director de herbario, una inspiracio6n como mentor para sus 
estudiantes y colegas, y un noble amigo. 


One of the most renowned American plant taxonomists of the 20th century 
has died. It is with great sadness and a profound sense of loss that this reflective 
tribute is written about the life and career of John W. Thieret, Professor Emeri- 
tus of Biological Sciences at Northern Kentucky University, retired Director of 
the Northern Kentucky University Herbarium, and Associate Editor of Sida, 
Contributions to Botany, and Editor of the Journal of the Kentucky Academy of 
Science JKAS). John suffered a brain aneurysm at his home in Alexandria, Ken- 
tucky, on 6 December 2005, while editing a manuscript for the JKAS. He never 
regained consciousness and died on 7 December. He was 79 years old. 

John William Thieret was born on | August 1926, in Chicago, Illinois, the 
only child of Hans and Lorena Thieret. Growing up, he was interested in plants 
and became an avid student of botany during his school days at Hyde Park High 
School. At Hyde Park, John met his future wife, Mildred Wolf, fittingly in a 
botany class. After working briefly in Chicago, John moved to Logan, Utah, to 
study at Utah State University. Three years later, Mildred also moved to Logan 
and attended Utah State. They were married on 13 March 1950, by one of their 
professors, after completing their Evolution final exams. Both earned their B.S. 
degrees in 1950: John’s in Botany and Mildred’s in Bacteriology. They remained 
at Utah State University for graduate work and in 1951, John earned his MS. in 
Botany with a thesis in barley genetics and Mildred completed her MS. in Bac- 
teriology. They then returned to Chicago, where John attended the University 
of Chicago to work on his doctorate under Theodor K. Just, Chief Curator of the 
Chicago Field Museum of Natural History. John received his Ph.D. in Botany in 


SIDA 22(1): 3-19. 2006 


4 BRIT.ORG/SIDA 22(1) 


John’s senior kink n I nhat tHy in Darl Hig! C-} |, Chicago, ca 1946. 


3 r 


1953. The title of John’s dissertation was “Gross Morphology of the Seeds of the 
Scrophulariaceae and Classification of the Family.” 


Later in 1953, John became Assistant Curator of Economic Botany at the 
Chicago Field Museum and then Curator of Economic Botany from 1954 to 1961. 
While at the Field Museum, he made collecting trips to Cuba, Mexico, the north- 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 5 


ern Great Plains of the United States, and the Northwest Territories of Canada. 
John authored 26 publications during that time and his special interests in the 
Poaceae and Scrophulariaceae were evident in his published works. John pub- 
lished five new nomenclatural combinations while at the Field Museum. He 
also wrote three articles on the flora and vegetation of the Canadian Northwest 
Termitories, 

John left the Field Museum to become Associate Professor and later Profes- 
sor of Biology at the University of Southwestern Louisiana, Lafayette (USL) from 
1961 to 1973. At USL, he sponsored undergraduate research projects, and directed 
six MS. theses and one Ph.D. dissertation. During this period, John conducted 
most of his personal research in the southeastern United States, with an em- 
phasis on the Louisiana flora. At USL, he authored or co-authored 47 articles. 
John named four plant species new to science that he discovered in Louisiana: 
Cyperus brevifolioides Thieret & Delahoussaye; Cyperus louisianensis Thieret; 
Isoetes louisianensis Thieret; and Limnophila x ludoviciana Thieret. From his 
work at USL, John published 11 nomenclatural combinations. He also published 
his first two generic flora treatments of the southeastern United States, five more 
articles from his investigations in the Canadian Northwest Territories, and sev- 
eral new United States or North American plant records. 

John made his final academic career move in 1973, when he joined the fac- 
ulty at Northern Kentucky University (NKU) in Highland Heights, as Profes- 
sor and Chair of the Department of Biological Sciences. The opportunity to teach 
at NKU, the presence of the Lloyd Library in Cincinnati, and relocating his fam- 
ily to live in a cooler climate, were especially appealing to him. John served as 
Chair until 1980 and continued as Professor until retiring in 1992, with the title 
Professor Emeritus of Biological Sciences. 

John loved teaching and was an exemplary teacher in the classroom, labo- 
ratory, and field. His classes were challenging, enjoyable, informative, popular, 
and he helped many students reach their full potential. He always was avail- 
able to students and colleagues for discussions about botany, academics, or just 
about life. John was a strong believer in providing students with actual plant 
specimens, either fresh or dried, for a “hands-on” approach, and he went to great 
effort to accomplish that goal. In the field, John’s passion for botany was espe- 
cially contagious and he always felt rejuvenated after a field trip with students. 
John generously shared his expertise and enthusiasm for botany, especially plant 
taxonomy, with students, colleagues, and the general public for 32 years. His 
exuberance, magnetic personality, and knowledge were inspirations to every- 
one his activities touched. 

For many years, John’s summer activities were focused on teaching at vari- 
ous biological field stations. He served as Visiting Lecturer in Botany at the Itasca 
Biological Station, University of Minnesota; the Oklahoma Biological Station, 
University of Oklahoma; the Michigan Biological Station, University of Michigan; 


— 


— 


6 BRIT.ORG/SIDA 22(1) 


faculty pt t University of Soutt Louisi Lafayette, 1972. 


and the Franz Theodore Stone Laboratory, Ohio State University. John enjoyed 
the flora of cooler climates, and biological stations provided Mildred and his 


children (Robert, Nancy, Richard, Jeffrey, and Jennifer) new environments to 
experience. He also conducted major field travels in the southeastern, south- 
western, and northwestern United States, the Great Basin of Nevada and Utah, 


and the Canadian Arctic, Newfoundland. and Ontario. 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 7 


- Bate. ve 


, ae 4 


a 
‘a 


be 5 


John with Fraser magnolia in West Virginia during 1998 (D.M. Brandenburg). 


John authored or co-authored with other botanists, 58 refereed journal ar- 
ticles at NKU. He frequently collaborated in research activities and publica- 
tions with current and former undergraduate and graduate students and 
younger NKU colleagues. Indeed, he was a mentor whose enthusiasm was con- 


8 BRIT.ORG/SIDA 22 


= 


1) 


tagious and inspired botanists from many other institutions to pursue research 
activities. The scope of his collaborative research ranged from investigations in 
Kentucky, research elsewhere in the United States, to the massive North America 
flora project. John was the prime mover and inspiration of the Kentucky flora 
project, which culminated with the publication of Ronald L. Jones’ Plant Life of 
Kentuckyin 2005. John served as Editorial Associate and contributed the treat- 
ment of the Poaceae for this comprehensive flora. 

During his tenure at NKU, John was the author or co-author of five books. 
These books were entitled: Louisiana Ferns and Fern Allies; Aquatic and Wet- 
and Plants of Kentucky; Trees: A Quick Reference Guide to Trees of North 
America; Assessment and Management of Plant Invasions, and National Audu- 
bon Society Field Guide to North American Wildflowers: Eastern Region. His 
boo 


— 


<s continue to serve many audiences, from professional botanists to ama- 
teur plant enthusiasts. 

One of John Thieret’s crowning achievements was establishing the North- 

ern Kentucky University Herbarium (KNkK) in 1973. As its first Director, he built 
the herbarium from his personal collections, additions by colleagues and stu- 
dents, and through an active specimen exchange with various national and 
international herbaria. Currently, KNK has over 35,000 mounted specimens 
with strong representation from Kentucky, the Southeast, and the Midwest. The 
KNK collection has the highest species diversity among Kentucky herbaria col- 
lections. Through his efforts, the KNK herbarium has become the best-curated 
herbarium in Kentucky. John donated his entire 600 book reference library to 
the herbarium to further support botanical research. 

John was a consummate field botanist whose excellent field abilities and 
extensive experiences were essential components of hisaccomplishments. He had 
an incredible memory forall of the plants that he had ever encountered and would 
make accurate field identifications even if he had seen the plants only once 
from several decades ago. This lifetime inventory combined with an amazing 
eye for detail enabled John to locate species that others would pass. John made an 
astounding number of new state, regional, United States, and North American 
plant records during his career. Most contemporary alpha plant taxonomists 
concentrate their study ona few groups, write floras at the local or state level, 
pursue floristics, or other taxonomic works. John was one of the last of a rare and 
remarkable group of field naturalists whose breadth and depth of knowledge 
of botanical life throughout the United States and Canada was unparalleled in 
his later years. He was an ardent plant collector throughout his professional 
career with specimen accession numbers over 62,000. John accurately identified 
all plants he deposited into various herbaria and meticulously mounted hisown 
plant specimens. 


— 


John Thieret was an internationally known botanist, excellent educator, 
wise scholar, and gentleman. He dearly loved his family, and after family, his 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 9 


Leet . 
UNIVETOILY, 


. 
7 
> 
5 


Ink ° ob NIWIT | 


, J 4 


10 BRIT.ORG/SIDA 22(1) 


great passion was plants. Although John was focused on plant taxonomy and 
learning new plants his entire life, he had other diverse interests including clas- 
sical music, particularly opera. He also enjoyed sharing his knowledge far be- 
yond botany and the natural sciences through verbal discourse about diverse 
topics in history, literature, art, religion, and the social sciences. 

Scientists often are judged by the number of articles by them or about them. 
If this is any measure of a person, John stands taller than a coastal redwood. Dur- 
ing his professional career, John authored at least 157 refereed journal articlesand 
book articles. Forty-two articles appeared in Sida, Contributions to Botany, which 
was his favorite journal. He also regularly published in Bartonia, Canadian Field- 
Naturalist, Castanea, Economic Botany, Journalofthe Arnold Arboretum, Rhodora, 
Taxon,and Transactions/Journalofthe Kentucky Academy of Science. In addition 
to books, journal articles, and 19 nomenclatural com binations, John wrote 136 
book reviews, 65 articles for Encyclopaedia Britannica, 46 articles for Encyclope- 
dia Americana, and many popular science articles. 

John served several significant editorships and advisory roles during his 
professional career. He wasa Member of the Editorial Board of Economic Botany 
from 1959-1965, Book Editor of Economic Botany from 1959-1984, Editor from 
1986-1990, and Associate Editor from 1992-2002. He was a founding Member 
of the Editorial Board for the Vascular Flora of the Southeastern United States 
project from 1981-2005, and wrote seven southeastern generic flora treatments. 

John served as the Associate Editor of Sida, Contributions to Botany, from 
1972-2005, and contributed to its excellence and prestige through his dedicated 
work. Barney L. Lipscomb, current Editor of Sida, Botanical Research Institute 
of Texas, notes, ‘John wasa towering lighthouse to the editors of Sida. His stead- 
fast editorial, ‘light, was a never ending source of guidance and navigation in 
keeping Sida on a positive course.” 

From 1981-1995, John served as Abstract Editor of Transactions of the Ken- 
tucky Academy of Science (TKAS). He then assumed the position of Editor of 
TKAS in 1996 and changed the title from Transactions to Journal of the Ken- 
tucky Academy of Science JKAS) in 1998 to better reflect its content. Through 
his efforts, John greatly enhanced the stature of the JKAS as a multi-disciplin- 
ary journal of state and regional scientific literature. 


During 1983-2005, one of his most significant roles was serving asa Mem- 
ber of the Editorial Committee of the monumental multi-volume project, Flora 
of North America North of Mexico. He edited the first 10 published volumes and 
prepared 25 family and generic treatments. John also was an Advisor in Botany 
tor Encyclopaedia Britannica from 1959-2005 and a Member of the Advisory 
Committee at the Lloyd Library in Cincinnati from 1992-2005. 

John loved editing and his ability as an editor was extraordinary. He was a 
perfectionist and a superlative editor second to none, but his efforts always 
brought out the best in authors. Those who submitted manuscripts for John’s 


em 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 11 


editorial scrutiny often found them returned with a profusion of red ink on the 
printed text. When his recommended changes were made, the greatly improved 
manuscripts always told the story better. 

John received many awards and honors during his career. One of his most 
esteemed honors was having a mint in the Lamiaceae that he discovered in Loui- 
siana named for him. In 1964, botanist Lloyd H. Shinners named this new spe- 
cies Scutellaria thieretii Shinners. 

He received the 1984 Distinguished Kentucky College/University Scien- 
tist Award from the Kentucky Academy of Science for his significant academic 
research and teaching contributions to the Commonwealth. John was recently 
presented the 2005 Outstanding Academy Service Award from the Kentucky 
Academy of Science for his outstanding editorial contributions to the JKAS. 

To commemorate his contributions to the Northern Kentucky University 
Herbarium, it was officially renamed The John W. Thieret Herbarium by the 
Northern Kentucky University Board of Regents on 22 March 2000. 

In 1994, he established The John W. Thieret Research Award as an annual 
award to the NKU biological sciences student who accomplished the most sig- 
nificant research. The Thieret family would like to continue this student honor, 
and have asked that donations in his memory be sent to the Northern Kentucky 
University Foundation, designated to that research award. 

John is survived by his devoted wife of 55 years, Mildred Thieret, his five 
children, Robert, Nancy, and Jeffrey in Minnesota, Richard in China, and Jenni- 
fer in Kentucky, seven grandchildren, and five great-grandchildren. 

John Thieret was one of the patriarchs of North American plant taxonomy, 
and one of the last great field naturalists of the 20th century. His death marks 
the ending of a botanical era, but his legacy continues through the work of many 
former students and colleagues. John was an inspiring and stabilizing mentor 
who enriched the lives of those who knew him. He will be greatly missed by all. 


ACKNOWLEDGMENTS 


lam especially grateful to Mildred Thieret for graciously sharing her memories 
about John and providing photographs of her husband. lalso acknowledge con- 
tributions from Barney L. Lipscomb, Botanical Research Institute of Texas, 
Ronald L. Jones, Eastern Kentucky University, David M. Brandenburg, Dawes 
Arboretum, James O. Luken, Coastal Carolina University, and Debra K. Pearce, 
Northern Kentucky University. 


REFEREED PUBLICATIONS 


Bairp, J.R.and J.W. THierer. 1985. Notes on Themeda quadivalvis (Poaceae) in Louisiana. |selya 
2:129-137. 

Bair, J.R.and J.W. THierer. 1988.The bur gherkin (Cucumis anguria var.anguria, Cucurbitaeae). 
Econ. Bot. 42:447-451. 


12 BRIT.ORG/SIDA 22(1) 


Bairo, J.R.and J.W. THierer. 1989. The medlar (Mespilus germanica, Rosaceae) from antiquity 
to obscurity. Econ. Bot. 43:328-372 

Baird, JR. and JW. Tuierer. 1993. Spartina. In: J.C. Hickman, ed. The Jepson manual: higher 
plants of California. University of California Press, Berkeley and Los Angeles. Pp. 1296- 
1297. 

Beat, E.O. and J.W. THieret. 1986. Aquatic and wetland plants of Kentucky, Kentucky State 
Nature Preserves Commission, Scientific and Technical Series, Number 5, Frankfort. 
BranbensurG, D.M., W.H. BLackwelt, and J.W. THiereT. 1991. Revision of the genus Cinna 

(Poaceae). Sida 14:581-596. 

Branoensure, D.M., J.E. Estes, $.B. Russet, and J.W. THieret. 1991. One-nerved paleas in Cinna 
arundinacea L. (Poaceae). Trans. Kentucky Acad. Sci. 52:94-96. 

BraNnbenBurG, D.M., J.E. Estes, and J.W. THierer. 1991.Hard grass (Sclerochloa dura, Poaceae) in 
the United States. Sida 14:369-376. 

BranbensurG, D.M. and J.W. THieret. 1996, Sclerochloa dura (Poaceae) in Kentucky. J. Ken- 
tucky Acad. Sci. 57:47-48. 

stp D.M. and J.W. THierer. 2000. Cinna and Limnodea (Poaceae): not congeneric. 
Sida 19:195-200. 

ree D.M. and J.W. THieret. 2003. Epipactis helleborine (Orchidaceae) in Kentucky, 
with overview of literature on biology of the species. J. Kentucky Acad. Sci. 64:55-74. 

Buopett Il, G.F.and J.W. THieret. 1985. Notes on Frigenia bulbosa (Apiaceae). Bartonia 51:69-76. 

Bupbett Il, G.F. and J.W. THieret. 1997. Saururaceae. In: Flora of North America Editorial 
Commitee, eds. Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford, 
Pp. 37-38 

Buooett II,G.F.and J.W. THierer. 2004.Lobed leaves in Salix exigua, sandbar willow (Salicaceae), 
in Kentucky. J. Kentucky Acad. Sci.65:51. 

Caraway, M.L. and J.W. THieret. 1985. Amphibromus scabrivalvis (Gramineae) in Louisiana. 
Sida 11:207-214. 

Crark, H.L. and J.W. THierer. 1968. The duckweeds of Minnesota. Michigan Bot. 7:67-76. 

Crark, R.C., R.L. Jones, TJ. Weckman, R.L. THomeson, J.W. THieReT, Kentucky State Nature Preserves 
Commission, and K. Feeman. 2005. State records and other noteworthy collections for Ken- 
tucky. Sida 21:1909-1916. 

CrANFILL, R. and J.W. Trieret. 1981. Thirty additions to the vascular flora of Kentucky. Sida 
9:55-58. 

DetaHoussaye, AJ, and J.W. THieret. 1967. Cyperus subgenus Kyllinga (Cyperaceae) in the 
continental United States. Sida 3:128-136. 

Eckenwatoer, J.E. and J.W. THieret. 1993. Keys to gymnosperm families. In: Flora of North 
America Editorial Committee, eds. Flora of North America, Vol. 2. Oxford Univ. Press, 
New York and Oxford. Pp. 345-346. 

Evers, R.A.and J.W. THieret. 1957.New plant records: Illinois and Indiana. Rhodora 59:181. 

Hawt, D.W.and J.W. THierer. 2003. Chrysopogon. in: Flora of North America Editorial Commit- 
tee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 
633-636. 

HARTMAN, R.L., J.W. THieret, and R.K. Raseter. 2005. Paronychia. |In:Flora of North America Edito- 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 13 


rial Committee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and 
Oxford. Pp. 30-43. 

Hits, M.H., J.W. THieret, and J.D. MorerieLo. 2003. Sarcobatus.|n: Flora of North America Edito- 
rial Committee, eds. Flora of North America, Vol. 4. Oxford Univ. Press, New York and 
Oxford. Pp. 387-389. 

Kartesz, J.T., P Atten, and J.W. THieret. 1997. Epilobium brachycarpum (Onagraceae) in Ken- 
tucky. Trans. Kentucky Acad. Sci. 58:99. 

Kartesz, J.T. and J.W. THieret. 1991. Common names for vascular plants: guidelines for use 
and application. Sida 14:421-434. 

Lanory, G.and J.W. THiereT. 1973./soetes louisianensis 
Siana. Sida 5:129-130. 

Luken, J.O. and J.W. THierer. 1987a. Linum grandiflorum (Linaceae), Papaver dubium 
(Papaveraceae), and Salvia pratensis (Labiatae): additions to the Kentucky flora. Trans. 
Kentucky Acad. Sci. 48:26. 

Luken, J.O.and J.W. Tuierer. 1987b.Sumac-directed patch succession on northern Kentucky 
roadside embankments. Trans. Kentucky Acad. Sci. 48:51—54. 

LukeN, J.O. and J.W. Tierer. 1988. A life-form spectrum for Ohio. Trans. Kentucky Acad. Sci. 
49:38-39. 

Luken, J.O. and J.W. THieret. 1995. Armur honeysuckle (Lonicera maackii; Caprifoliaceae): its 
ascent, decline, and fall. Sida 16:479-503. 

Luxen, J.O.and J.W. THierer. 1996. Amur honeysuckle, its fall from grace. BioScience 46:18—24. 

LuKen, J.O. and J.W. THieret. 1997. Amur honeysuckle, its fall from grace. Arnoldia 57:2-12. 

Luken, J.O. and J.W. THigret. 2001. Floristic comparisons of mud flats and shorelines at Cave 
Run Lake, Kentucky. Castanea 66:336-351. 

Luken, J.O. and J.W. THieret (eds.). 1997. Assessment and management of plant invasions. 
Springer-Verlag, Inc., New York, NY. 

Luken, J.O., J.W. THieret, and J.R. Karresz. 1993. Erucastrum gallicum (Brassicaceae): invasion 
and spread in North America. Sida 15:569-582. 

Meotey, M.E., H. Bayan, J. MacGrecor, and J.W. THieret. 1985.Achyranthes japonica (Miq.) Nakai 
(Amaranthaceae) in Kentucky and West Virginia:new to North America. Sida 11:92-95. 

Meotey, M.E., R. Craneitt, and J.W. THieret. 1983. Vascular flora of Kentucky: additions and 
other noteworthy collections. Sida 10:114-122. 

Meotey, M.E. and J.W. THierer. 1991. Ulmus parvifolia (Ulmaceae) naturalized in Kentucky. 
Sida 14:610-613. 

MoHtensrock, R.H. and J.W. THieret. 1987. Trees: a quick reference guide to trees of North 
America. Collier Books, Macmillan Publishing Company, New York, NY. 

Naczi, R.F.C. and J.W. THieret. 1996a. Addition to the flora of Bradford County, Pennsylvania. 
Bartonia 59:81—-85. 

Naczi,R.F.C.and J.W. THieret. 1996b.Invasion and spread of Coincya monensis (Brassicaceae) 
in North America. Sida 17:43-53. 

Naczi, R.F.C. and J.W. THieret. 1996c. The gold-cone tamarack (Larix larcina forma lutea, 
Pinaceae) in Pennsylvania. Bartonia 59:123-124. 


Isoetaceae), a new species from Loui- 


— 


14 BRIT.ORG/SIDA 22(1) 


Naczi, R.F.C. and J.W. Tuierer. 2000. Additions to the flora of Potter County, Pennsylvania. 
Bartonia 60:117-120. 

Nienaser, M.A. and J.W. Tuieret. 2003. Phytolaccaceae. In: Flora of North America Editorial 
oo eds. Flora of North America, Vol.4.Oxford Univ. Press, New York and Oxford. 
Po 

Pearce, D. . eae JW. Tuieret. 1991. Japanese-quince (Chaenomeles speciosa, Rosaceae), a 
dual-use shrub. Econ. Bot. 45:285-288. 

Pearce, D.K.and J.W. Tuieret. 1993. Persimmon (Diospyros virginiana, Ebenaceae) and may- 
apple (Podophyllum peltatum), Berberidaceae): proximate analysis of their fruits. Trans. 
Kentucky Acad. Sci. 54:30-31. 

Raseter, R.K.and J.W. THieret. 1988.Comments on the Caryophyllaceae of the southeastern 
United States. Sida 13:149-156. 

Raseter, R.K.and J.W.THierer. 1997. Sagina (Caryophyllaceae) range extensions in Canada:S. 
japonica new to Newfoundland, S.procumbens, new to the Northwest Territories. Canad. 
Field-Naturalist 111:309-310. 

Reece, W.D. and J.W. Tuieret. 1966. Botanical study of the Five Islands of Louisiana. Castanea 
31:251-277. 

Smith, C.E, Jr.and J.W. THieret. 1959a.An English obituary account of Thomas Nuttall. Bartonia 


—s 


Smith, C.E., Jr.and J.W. THieret. 1959b. Thomas Nuttall (1786-1859): an evaluation and bibli- 
ography. Leafl. West. Bot. 9:33-42. 
SpactH, J.P. and J.W. THieret. 2004. Notes on “coffee” from the Kentucky coffeetree 
(Gymnocladus dioicus, Fabaceae). Sida 21:345-356. 
Spuut, R.W. and J.W. Tuierer. 1989. Confusion between multiple and aggregate fruits. Bot. 
Rev. 55:53-72. 
Tuieret, J.W. 1953. Gross morphology of the seeds of the Scrophulariaceae and classifica- 
tion of the family. Ph.D. dissertation. The University of Chicago. 
THieret, J.W. 1953. Dipsacus laciniatus in Illinois. Rhodora 55:268. 
THiereT, J.W. 1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4: 
164-184. 
THieret, J.W. 1955a. The seeds of Veronica and allied genera. Lloydia 18:37-45. 
THiereT, J.W. 1955b. The status of Berendtiella A. Gray. Ceiba 4:304—-305. 
THieret, JW. 1956a. Bryophytes as economic plants. Econ. Bot. 10:/5-91. 
THieReT, JW. 1956b. Nardoo. Amer. Fern J. 46:108-109. 
] 
] 
] 


— 


THieret, J.W. 1956c. Stenardrium Nees versus Gerardia L. Taxon 5:58-59. 

THiereT, J.W. 1957. Plants new to Illinois and to the Chicago region. Rhodora 59:289. 

Tuieret, J.W. 1958a.Agalinis Rafinesque versus Chytra Gaertn. Taxon 7:142-143. 

THieret, J.W. 1958b. Castilleia Mutis ex L. versus Bartsia L. Taxon 7:83-84. 

THiereT, J.W. 1958c. Economic botany of the cycads. Econ. Bot. 12:3-41. 

Tuieret, J.W.1959a. Grassland vegetation near Fort Providence, Northwest Territories. Canad. 
Field-Naturalist 73:161-167 

TuiereT, J.W. 1959b, Scrophulariaceae. In: J. Angely (ed). Catalogo e estatastica dos géneros 
botanicos fanerogamicos, Vol. 49. 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 15 


THiereT, J.W. 1960a. Calamovilfa longifolia and its variety magna. Amer. Midl. Nat. 63: 
169-176. 

THiereT, J.W. 1960b.The formaldehyde method of collecting plant specimens. Turtox News 
38:114-115 

THieret, J.W. 1961a. A collection of plants from the Horn Plateau, District of Mackenzie, 
Northwest Territories. Canad. Field-Naturalist 75:77-83. 

THierET, J.W.1961b.New plant records for southwestern District of Mackenzie.Canad. Field- 
Naturalist 75:111-121. 

THieret, JW. 1961¢. The Scrophulariaceae-Buchnerae of Central America. Ceiba 8:92-101. 

THiereT, J.W. 1961d.The specific epithet of the pecan. Rhodora 63:296. 

TuiereT, J.W. 1962a. Exceptional height for Rhododendron lapponicum. Canad. Field-Natu- 
ralist 76:123. 

Tiere, J.W. 1962b. New plant records from District of Mackenzie, Northwest Territories. 
Canad. Field-Naturalist 76:206-208. 

TuiereT, J.W. 1963a. Additions to the flora of Louisiana. Castanea 28:169-1 70. 

TuiereT, J.W. 1963b. Additions to the flora of the Northwest Territories. Canad. Field-Natu- 
ralist 77:126. 

Tuieret, J.W. 1963c. Botanical survey along the Yellowknife Highway, Northwest Territories, 
Canada. |: Catalogue of the flora. Sida 1:117-170 

THierET, J.W. 1963d. Life-forms in the plains flora of southern Mackenzie, Northwest Territo- 
ries. Rhodora 65:149-157. 

TuiereT, J.W. 1963e. The correct name for the watermelon. Taxon 12:37. 

Tieret, J.W. 1964a. Botanical survey along the Yellowknife Highway, Northwest Territories, 
Canada. II: Vegetation. Sida 1:187-239. 

TuiereT, J.W. 1964b. Eriogonum annuum (Polygonaceae): biennial in Nebraska. Sida 1:382. 

Tuieret, J.W. 1964c. Fatoua villosa (Moraceae) in Louisiana: new to North America. Sida 
1:248. 

Tuieret, J.W. 1964d. Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) 
in Louisiana: new to the United States. Sida 1:249-250. 

TuiereT, J.W. 1964e. More additions to the Louisiana flora. Sida 1:294-295. 

TuHiereT, J.W. 1966a. Additions to the Louisiana flora. Sida 2:264—-265. 

THiereT, J.W. 1966b. Habit variation in Myrica pensylvanica and M. cerifera. Castanea 31: 
183-184. 

THiereT, J.W. 1966c. Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2: 


THiereT, J.W. 1966d. Synopsis of the genus Calamovilfa (Gramineae). Castanea 31:145-152. 

THieRET, J.W. 1967a.Life-forms in the flora of Minnesota. J. Minnesota Acad. Sci.34:251-277. 

Tuierer, J.W. 1967b. Neogaerrhinum kelloggii (Greene) Tuieret, comb. nov. (Scrophulariaceae) 
Sida 3:187, 

THieret, J.W. 1967c. Supraspecific classification in the Scrophulariaceae: a review. Sida 3: 
87-106. 

Tuieret, J.W. 1967d. Thirty additions to the Louisiana flora. Sida 3:123-127. 


BRIT.ORG/SIDA 22(1) 


THierET, J.W. 1968. Additions to the vascular flora of Louisiana. Proc. Louisiana Acad. Sci. 
31:91-97 
THieret, J.W.1969a. Baptisia lactea (Rafinesque) THieret, comb.nov. (Leguminosae). Sida 3:446. 
THiERET, J.W. 1969b. Notes on Epifagus. Castanea 34:397-402, 
THiereT, J.W. 1969c. Rumex obovatus and Rumex paraguayensis (Polygonaceae) in Louisi- 
ana: new to North America. Sida 3:445-446. 
THieret, J.W. 1969d. Sagittaria guayanensis (Alismataceae) in Louisiana: new to the United 
States. Sida 3:445. 
Tiere, J.W. 1969e. Trifolium vesiculosum (Leguminosae) in Mississippi and Louisiana: new 
to North America. Sida 3:446-447, 
THiereT, J.W. 1969f. Twenty-five species of vascular plants new to Louisiana. Proc. Louisiana 
Acad. Sci. 32:78-82. 
THiereT, J.W. 1970a. Bacopa repens (Scrophulariaceae) in the conterminous United States. 
Castanea 35:132-136. 
THIERET, J.W. 1970b. Nemophila microcalyx, an incorrect name. Rhodora 79:399-400. 
TuiereT, J.W. 1970c. Orobanchaceae. In: C.L. Lundell, ed. Flora of Texas, Vol. 2. Pp. 331-337. 
THiereT, JW. 1970d. Sere Df yhulariac eae—fiqwort family.In:E.A Menninger. ed Flowering vines 
of the world. Pp. 316-321. 
THIERET, J.W. 1971a. Additions to the Louisiana flora. Castanea 36:219-222. 
THiereT, J.W.1971b. Eriocaulon cinereum R.Br. in Louisiana. Southw. Nat. 15:391. 
THiereT, J.W.1971¢.Observations on some aquatic plants in northwestern Minnesota. Michi- 
gan Bot. 10:117-124. 
THierer, J.W. 1971d. Physalis lagascae (Solanaceae) in Louisiana: new to the conterminous 
United States. Sida 4:277. 
THiereT, J.W. 1971e. Quadrat study of a bottomland forest in St. Martin Parish, Louisiana. 
Castanea 36:174-181. 
THieret, JW. 1971f. The genera of Orobanchaceae in the southeastern United States. J. 
Arnold Arbor. 52:404-434. 
THieret, JW. 1972a. Aquatic and marsh plants of Louisiana: a checklist. J. Louisiana Soc. 
Hort. Res. 13:1-45. : 
THiereT, J.W.1972b. Checklist of the vascular flora of Louisiana: Part 1. Fern and fern allies, 
gymnosperms, and monocotyledons. Lafayette Nat. Hist. Mus, Techn. Bull. 2. 
THierET, J.W. 1972c. Rotala indica (Lythraceae) in Louisiana. Sida 5:45. 
THiereT, JW. 1972d. Synopsis of Hemichaena, including Berendtiella (Scrophulariaceae) 
Fieldiana Bot. 34:89-99, 
THieret, JW. 1972e. The Phrymaceae in the southeastern United States. J. Arnold Arbor. 
53:226-233. 
THieRET, J.W. 1972f. Zeuxine strateumatica in Louisiana. Amer. Orch. Soc, Bull. 41:413. 
THiereT, J.W. 1973. Sex and the angiosperms. Sida 5:59-60. 
THieret, J.W.1974.Allium ampeloprasum (Liliaceae) and Trifolium vesiculosum (Leguminosae) 
in Oklahoma. Sida 5:286-287. 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 17 


THieret, JW. 1975a. Hemigraphis reptans (Acanthaceae), a greenhouse weed in Louisiana. 
Sida 6:115. 

THiereT, J.W. 1975b. The Mayacaceae in the southeastern United States. J. Arnold Arbor. 
56:248-255. 

THiereT, J.W. 1976a. Floral biology of Proboscidea louisianica (Martyniaceae). Rhodora 78: 
169-179. 

TuiereT, J.W. 1976b. Vascular plants new to Ohio. Castanea 41:181-183. 

THieret, J.W.1977a. Cyperus louisianensis (Cyperaceae), a new species from southern Loui- 
Siana. Proc. Louisiana Acad. Sci. 40:23-26. 

THiereT, J.W.1977b. Juvenile leaves in Oklanoma Marsilea (Marsileaceae). Sida 7:218-219. 

THieret, J.W. 1977c. Life-forms in the Michigan flora. Michigan Bot. 16:27-33. 

THiereT, J.W. 1977d. The Martyniaceae in the southeastern United States. J. Arnold Arbor. 


THieRET, J.W. 1979. Hyptis mutabilis (Labiatae) in southeastern United States. Sida 8: 

202-209. 

TiereT, J.W. 1980. Louisiana ferns and fern allies. Lafayette Natural History Museum. Pub- 
lished in conjunction with The University of Southwestern Louisiana, Lafayette. 

TuiereT, J.W. 1982. The Sparganiaceae in the southeastern United States. J. Arnold Arbor. 
63:341-355. 

THierET, J.W. 1988. The Juncaginaceae in the southeastern United States. J. Arnold Arbor. 
69:1-23. 

TuierET, J.W. 1989. Picea abies (Pinaceae) naturalized in southeastern Minnesota. Sida 13:505. 

THieRET, J.W. 1993a. Calocedrus. In: Flora of North America Editorial Committee, eds. Flora of 
North America, Vol. 2. Oxford Univ. Press, New York and Oxford. P. 412. 

TuHiereT, J.W. 1993b. Pinaceae. In: Flora of North America Editorial Committee, eds. Flora of 
North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Pp. 352-354. 

THIERET, J.W.1993c. Psilotaceae. In: Flora of North America Editorial Committee, eds. Flora of 
North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Pp. 16-17. 

THieRET, J.W. 2003a. Arthraxon. In: Flora of North America Editorial Committee, eds. Flora of 
North America, Vol. 25.Oxford Univ. Press, New York and Oxford. P.677. 

TuHieret, J.W. 2003b. Calamovilfa. In: Flora of North America Editorial Committee, eds. Flora 
of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 140-144. 

Tuieret, J.W. 2003c. Coix.In: Flora of North America Editorial Committee, eds. Flora of North 
America, Vol. 25.Oxford Univ. Press, New York and Oxford. Pp. 703-704. 

THiereT, JW. 2003d. Eremochioa. In: Flora of North America Editorial Committee, eds. Flora 
of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 688-690. 

THierET, J.W. 2003e. Hackelochloa. In: Flora of North America Editorial Committee, eds. Flora 
of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp.691-693. 

THieReT, J.W. 2003f. Microstegium. In: Flora of North America Editorial Committee, eds. Flora 
of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp.623-624. 

THierer, JW. 2003g. Monoanthochloé. |n: Flora of North America Editorial Committee, eds. 
Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 28-30. 


18 BRIT.ORG/SIDA 22(1) 


TuiereT, J.W. 2005. Agrostemma. Pp. 214-215 In: Flora of North America Editorial Commit- 
tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. 

Tuieret, J.W. and C.M. Atten. 1974. Setaria pallide-fusca (Gramineae) in Louisiana. Castanea 
39:290-291. 

THieret, J.W. and J.R. Bairo. 1985. Thalaspi alliaceum (Cruciferae) in Kentucky and Indiana. 
Trans. Kentucky Acad. Sci.46:143-145. 

Tieret, J.W. and D.M. Branbensure. 1986. Scaevola (Goodeniaceae) in southeastern United 
States. Sida 11:445-453. 

Tuieret, J.W. and R.A. Evers. 1957. Notes on Illinois grasses. Rhodora 59:123-124. 

Tiere, J.W.and S.F.Gtassman. 1958. Grasses new to Illinois and the Chicago Region.Rhodora 
60:264. 

THiereT, J.W., R.L. HARTMAN, and R.K. Rasecer. 2005. Herniaria. In: Flora of North America Edito- 
rial Committee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and 
Oxford. Pp. 43-45. 

THieret, J.W.and J.T. Kartesz. 1997. Lardizabalaceae.In:Flora of North America Editorial Com- 
mittee, eds. Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford. Pp. 
293-294. 

THiereT, J.W. and B.L. Lipscome. 1985. Scaevola sericea Vahl var. taccada (Gaertn.) THiereT and 
Lipscomb. Sida 11:103. 

Tuieret, J.W.and J.O. Luxen. 1996. The Typhaceae in the southeastern United States. Harvard 
Pap. Bot.8:27-56. 

THiereT, J.W. and R.S. Maptes. 1979. Thelypteris interrupta (Polypodiaceae) new to Louisiana. 
Iselya 1:55, 

Tuieret, J.W.,W.A. Nierinc, and N.C. Oumsteao. 2001. National Audubon Society Field Guide to 
North American wildflowers: eastern region, revised edition. National Audubon Soci- 
ety, Alfred A. Knopf, Inc., New York, NY. 

THieRET, J.W. and R.K. Raseter. 2005a. Corrigiola. In: Flora of North America Editorial Commit- 
tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 
48-49, 

THiereT, J.W. and R.K. Rasever. 2005b. Polycarpaea. In: Flora of North America Editorial Com- 
mittee, eds. Flora of North America, Vol. 5.Oxford Univ. Press, New York and Oxford. Pp. 
23-25. 

THieret, J.W.and R.K. Rasever. 2005c. Polycarpon.|n: Flora of North America Editorial Commit- 
tee, eds. Flora of North America, Vol.5.Oxford Univ. Press, New York and Oxford. Pp. 25-26. 

THiereT, J.W.and R.K. Raseter. 2005d. Saponaria.In: Flora of North America Editorial Commit- 
tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 
157-158. 

Tuieret, JW. and R.K Raseter. 2005e. Scleranthus. In: Flora of North America Editorial Com- 
mittee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 
149-151. 

THiereT, J.W.and R.K. Ragecer. 2005f. Vaccaria.|n:Flora of North America Editorial Committee, 
eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. P. 156. 


THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 19 


Tiere, J.W.and R.L. THompson. 1984. Cleome ornithopodioides (Capparaceae):adventive and 
spreading in North America. Bartonia 50:25-26. 

THieret, J.W. and S.B. Younc. 1988. The Kerguelen-cabbage, Pringlea antiscorbutica 
(Brassicaceae). Econ. Bot. 42:288-291. 

THompson, R.L.and J.W. THieret. 1986.Alopecuru dinaceus (Poaceae) established in Ken- 
tucky. Trans. Kentucky Acad. Sci.47:138. 

Vincent, M.A. and J.W. THieret. 1987. Thymelaea passerina (Thymelaeaceae) in Ohio. Sida 
12:51-54. 

Wesset, M.V. and J.W. THierer. 2000. Agrimonia (Rosaceae) in Kentucky with notes on the 
genus. J. Kentucky Acad. Sci.61:146-162. 

Woopwaro, R.W. and J.W. THieret.1953.A genetic study of complementary genes for purple 
lemma, palea, and pericarp in barley (Hordeum vulgare L.). J. Agronomy 45:182-185. 


POPULAR SCIENCE ARTICLES 


Branbensure, D.W. and J.W. THierer. 1999. Phytological peregrinations in West Virginia. The 
Dawes Arboretum Newsletter 33(1):2-3. 

BranbensurG, D.W.and J.W. THieret. 2000. Hobnobbing with the Egyptians. The Dawes Arbo- 
retum Newsletter 34(10):3. 

BranbdensurG, D.W. and J.W. THieret. 2001. Rafinesque and us. Lloydiana 6(1):4—-9. Reprinted 
with photographs in: Daweswood 1(2):20-28, 2004. 

Buppett, Il, G.F. and J.W. THieret. 1981. Squawroot: it toils not. Explorer 23(2):28-29. 

Buooett, Il, G.F. and J.W. THieret. 1981. Harbinger-of-spring. Explorer. 24(1):24—26. 

Tiere, J.W. 1955. Dragon's blood. Nature Mag. 48(7):372-374, 388. 

TuiereT, J.W. 1974. The papaw, Ohio's “tropical” fruit. Explorer 16(3):17-19. 

TuiereT, J.W. 1975. Long live the seeds. Explorer 17(1):4-7. 

Tiere, J.W. 1975. Beechdrops—benign parasite. Explorer 16(3):12-14. 

THierET, J.W. 1977. Consider the foxglove. Explorer 19(1):4—7. 

THieret, J.W. 1978. Red cedar. Explorer 20(4):26-29. 

Tiere, J.W. and D.R. Bezanson. 1983. Dutchman’s-breeches and squirrel-corn. Explorer 
25(1):8-13. 

Tiere, J.W. 1996. Incredible edibles: the ginkgo. Lloydiana 1(1):14-16. 

Tiere, J.W. 1996. Incredible edibles: lupine seeds. Lloydiana 1(3):9-12., 

THiereT, J.W. 1997. Frankincense and myrrh. Lloydiana 1(4):7-9. 

Tiere, J.W. 1997. Incredible edibles: durian. Lloydiana 2(2):12-17. 

Tuieret, JW. 1997. Incredible edibles: insects. Lloydiana 2(4):9-12. 

THiereT, J.W. 1998. Incredible edibles: teff. Lloydiana 3(2):4-7. 

THiereT, J.W. 1999. Incredible edibles: corn smut. Lloydiana 4(2):12-16. 

Tuieret, J.W. 1999. On the trail of Rafinesque. Flora of North America 13(3-4):13. 

THiereT, J.W. 2005. Kentucky's tropical fruit, the papaw. The Lady-Slipper 20(3):2-3. 

Ware, T.R.and J.W. THierer. 1987. Boomer of the prairie. Explorer 29(1):7-10. 


pay 


JOHN W. THIERET, THE CURATOR 


Robert F.C. Naczi 
Claude E. Phillips Herbariur 
Department aan & nie ee 
Delaware State University 
pee Se 9901, U.S.A. 
raczi@desu.edu 


John Thieret’s complex and remarkable personality hada way of leaving its mark 
on all who encountered him; those who met John and came to know him even 
briefly have rich and lasting memories of him. 1am the one who, in many ways, 
had to step into John’s shoes soon after his retirement from Northern Kentucky 
University, where | began my academic career in 1992. Of course, I could never 
fill John’s shoes, and I soon learned to tell my new-found NKU colleagues on 
first meeting them that | was occupying the position John had vacated, but | 
was not John’s “replacement.” Most of the time, I received knowing chuckles in 
response to such a self-introduction. 

Occupying the academic appointment John had held also meant I was the 
first toassume a curatorship for the herbarium that John built. After his arrival 
at NKU in 1973 as Chairman of the Biology Department, John lost no time in 
establishing a herbarium there. During my tenure as curator at NKU, John re- 
mained very active—in effect, John and I served as co-curators of the herbarium 
for nine years, to my great benefit. This relationship makes me uniquely quali- 
fied to remember John The Curator, a side of John to which | will limit the rest 
of my remarks. 

John strove for excellence in everything concerning the NKU Herbarium. 
He put his heart and soul into collecting, identifying, labeling, mounting, and 
filing plant specimens in the herbarium. Though he collected far more speci- 
mens than most botanists (in the several tens of thousands), he was never in 
competition with others to beat their collecting records. Rather, he was inter- 
ested, above all, in quality over quantity. Instead of complaining about the te- 
dium of pressing plants, typing labels, or gluing specimens on mounting paper, 
John took sincere pleasure in executing these tasks that many field botanists 
foist on assistants. Once a specimen was in the herbarium, he insisted on the 
highest standards for its curation. Many times when I hosted visitors to the NKU 
Herbarium, or | traveled to use other herbaria in Kentucky, | heard the remark 
that the NKU Herbarium was the best curated herbarium in the state of Ken- 
tucky. Within a short time, | came to see for myself that this statement was true. 

John had a photographic memory, which served him very well as curator. | 
can best illustrate John’s memory by relating an experience that continues to 


SIDA 22(1): 21 — 23. 2006 


22 BRIT.ORG/SIDA 22(1) 


amaze me. In the late spring of 1996, John entered the herbarium the day after I 
had returned from a long field trip through the southeastern United States for 
my research. John found me puzzling over a specimen I had collected. Though 
I was familiar with most flowering plants present in the springtime in decidu- 
ous forests of the Southeast, | had found something in one Alabama forest that 
completely stumped me. | had never seen this species, didn’t know its family, 
and couldn’t even be sure it was a monocot, though | somehow thought so. Turn- 
ing to John, | admitted that | didn’t know where to begin with this one, but 
suspected it was something “really good.” After all, what else could I say to cloak 
my ignorance? John gazed at my find, hesitated only a moment, didn’t say a 
word, and marched to one of several bookcases present in the herbarium. There, 
he removed one issue of one of the many scientific journals for which we had 
long series. Soon, he produced a photograph, and asked me, “Is this your plant?” 
With that photograph (Whetstone 1984: 133), I realized 1 had found the rare 
Croomia pauciflora(Nutt.) Torr. of the obscure family Stemonaceae. In response 
to my query if he had seen the plant, John responded, “No, but I remembered 
seeing that photograph appear in this paper many years ago.” 

When it came to desiderata for the herbarium, John was utterly ecumeni- 
cal. Whether the lowliest, most inconspicuous, most downtrodden weed or the 
loftiest tree, John was interested in learning its name, how to identify it, whether 
it was poisonous, and if he had any specimens of it in the herbarium. His en- 
thusiasm was infectious and quickly endeared him to other naturalists, par- 
ticularly in the field. When he encountered a plant species he hadn't seen in 
several years, he would treat the plant as if he’d run into a long-lost friend. My 
most recent field trip with John was last April, when he came to Delaware fora 
brief visit. linsisted on taking him toa nearby racetrack that was full of vernal 
weeds in all their unmown glory. Within minutes, John showed me Sibara 
virginica (L.) Rollins, a mustard species he knew well from Kentucky. Further 
investigation showed it to be previously undocumented for Delaware, though 
botanists have been hunting green treasure in this state since the earliest 1800s. 
Such was all in a day’s work if your combination of vast knowledge, keen eye, 
and unclouded memory were what made John Thieret the unparalleled botani- 
cal talent he was. 


— 


— 


John was very generous with imparting knowledge it had taken him years 
to amass. From him, | learned a tremendous amount, and have benefited im- 
mensely. Though John could be demanding, uncompromising, and prickly at 
times, he was also uncommonly human. So often when | expressed frustration 
at ill-prepared students, particularly ineffectual bureaucracy, or the insuffi- 
ciency of a 24-hour day, John would sigh and remark, “In a hundred years, no 
one will care a thing about it.” Yet, he cared about curation, cared about her- 
baria, and cared especially deeply for the NKU Herbarium, one of John Thieret’s 
great legacies. Hopelully, in a hundred years, people will care about John. 


NACZI, JOHN W. THIERET, THE CURATOR 23 


John Thi llecting herbari i Elko County, Nevada, 11 August 1993. 


REFERENCE 


WuetsTone, R.D. 1984. Notes on Croomia pauciflora (Sstemonaceae). Rhodora 86:131-137. 


JOHN W. THIERET, 
A STUDENT’S PERSPECTIVE (1967-2005) 


Robert R. Haynes' 
Department of Biological Sciences 
Box 870345 
Tuscaloosa, Alabama 35487-0345, U.S.A. 


It was a sad day 7 December 2005 when I was going through my e-mail and 
had two from Ron Jones. The first one indicated that John Thieret had suffered 
a massive brain hemorrhage, and the second one indicated that he had passed 
away. limmediately called Mrs. Thieret to offer her my condolences. I pointed 
out that I am now retired and live in rural North Louisiana. Consequently, | 
check my e-mail less often than while I was still a professor. Thus | had just 
learned of his illness. That was, by the way, the day he had passed. 

My first time to meet John Thieret was March 1967. I was in my final se- 
mester at Louisiana Tech, and he and | had agreed for me to attend The Univer- 
sity of Southwestern Louisiana (now University of Louisiana at Lafayette), where 
he was then employed, and to study under his direction for a master’s degree in 
plant systematics. Dr. Thieret at the time had a grant from NSF to prepare a 
Flora of Louisiana. Howard Clark, a graduate student at the time, and Dr. Thieret 
came to Louisiana Tech and took me on a weekend field trip to Northeastern 
Louisiana. We met about 6:00 a.m. at Carson Taylor Hall, the building that 
housed the Botany and Microbiology Department. Dr. Thieret had a truck with 
a camper that would sleep three people, which he had purchased with the NSF 
monies. In the rear of the camper were a few plants of flowering Sanguinaria 
canadensis (Papaveraceae), which they had collected the previous afternoon. 
That was my first time to see the species other than by herbarium specimen. 

During our weekend field trip, we visited quite a few beautiful deciduous 
forests, most with spring wildflowers. In particular we found Trillium (Liliaceae) 
(I cannot remember the species, and | am sure that John Freeman has since 
named that one diff ly than what Thieret would have called it). Again, that 
was my first experience with Trillium. The manner we used for locating good 
collecting habitat was to examine topographic maps and find areas with con- 
siderable relief. | thought at the time that we were quite fortunate in locating 
habitat. Since then I have learned much about field work, and | know that an 
excellent field botanist, such as John Thieret, has a sense for determining from 
topographic maps where good habitat might be. 

I began to learn the personality of John Thieret that weekend. He was con- 


Present Address: PO. Box 146, Athens, Louisiana 71003, U.S.A. 


SIDA 22(1): 25 — 31. 2006 


BRIT.ORG/SIDA 22(1) 


stantly smiling, making jokes, telling stories, and teaching. From that weekend 
to the day he died, he was always teaching whenever | was in his presence. 
George Fisher, who was a graduate student at Tech in 1967, and | went to 
Lafayette later that spring for a collecting trip to the Florida parishes with 
Thieret. | realized more about his practical jokes early on that trip. We followed 
his directions to the USL campus and Biology Building where he was awaiting 
our arrival. He asked us to follow him to his house where his cam per/truck was 
parked. He started home and took us through more back roads (all dirt) than 
one can imagine. When we arrived at his house (on a paved road of course), he 
got out of his car laughing saying he just wanted to have some fun. By following 
the paved road, one could get from the USL campus to his house in half the 


a 


time it took us! 

The Florida parish trip had a profound effect on my botanical career. In 
western Washington Parish, we collected a pondweed that Thieret called 
Potamogeton nodosus (Potamogetonaceae). | did not question him at all. During 
my first semester at USL, Dr. Thieret brought out the specimen and asked me to 
determine it. | said you called it P nodosus. He said that now he was not certain 
about that determination. I keyed out the specimen and consistently arrived at 
Potamogeton epihydrus, which was not supposed to be in Louisiana. I looked at 
the distribution, and it was known no closer than North Carolina at the time. | 
went to him, and he said that is what he determined the specimen to be also. 
That began a lifelong study of the genus Potamogeton. 

The fall of 1967 entered the graduate program at USL. Dr. Thieret had three 
other graduate students at the time, Howard Clark, Alex Lasseigne,and Billy Dan 
Hinton. Howard was working on the Lemnaceae of Louisiana, Alex was study- 
ing the Fabaceae (we called it Leguminosae at the time) of Louisiana, and Billy 
Dan was studying Parietaria (Urticaceae). | have always been interested in 
aquatic vascular plants, and my plans were to undertake a study of Utricularia 
(Lentibulariaceae) of Louisiana. Dr. Thieret began going through his literature 
and correspondence and learned that Peter Taylor of Kew had undertaken a 
study of Utricularia of North America and he, Dr. Thieret, did not want me to 
duplicate any study underway. Therefore, my future in the Lentibulariaceae 
came to an abrupt end. We talked about several genera, none of which really 
interested me, and we finally settled on Conopholis (Orobanchaceae), a genus 
as far from the aquatic environment as possible. As everyone who knows me is 
aware, I never really left the aquatic environment, however. 

Dr. Thieret had all of his graduate students work in the herbarium, and, 
current graduate students would find this hard to believe, without pay. We 
mounted and filed specimens, pulled specimens for loans, etc. As it turned out, 
| became the unofficial collections manager. | had closer interactions with Dr. 
Thieret than others for that reason. Of course, lalways received the blunt of any 
rage if things did not go just right, even if it was totally out of my control. 


a 


HAYNES, JOHN W. THIERET, A STUDENT’S PERSPECTIVE 27 


John Thieret loved classical music! He had a phonograph (tapes and CD's 
did not exist then!) that played in the herbarium much of the time, and he often 
placed it on one of the herbarium carts and played it in class. Thieret could 
whistle quite loudly, and he often was going around the biology building whis- 
tling classical music. 

Thieret suffered from occasional migraines, and, as lam sure most people 
with migraines, he was quite moody when he had a migraine. Whenever he 
was having a mood swing asa result of a migraine, he whistled exceptionally 
loudly. We graduate students learned to get out of the way whenever we heard 
that extremely loud whistling. In fact, if | was unusually late arriving in the 
morning and heard that whistle, I just went right on out of the building to a 
grill that was just off campus. All other graduate students had learned the rou- 
tine, and we all met at the grill for coffee until we thought the whistling had 
ceased. (I told him about our coffee excursions during a trip he and I made out 
West in 1998. He was totally unaware of these excursions, or so he said.) 

Without question, John Thieret was an editor, an excellent editor. I never 
will forget the first draft of my thesis. 1 proudly gave it to him, and it soon re- 
turned looking as if a chicken had stepped into red ink and then walked all 
over my manuscript! I learned from him to be much more careful when writ- 
ing and speaking. He had many favorite sayings, but one that | remember very 
well was related to editing. It is “consistency is next to godliness.” He said, be 
consistent; if you are incorrect at least be consistently incorrect! 

[learned much botany from John Thieret, but one thing in particular was 
that he did not like the term “pistil.” He said that it was an ambiguous term and 
that carpel was a much better term. | adopted his argument and never used or 
taught pistil after that. In courses that I taught, | always had to explain to the 
students why I did not use the term pistil regardless that it was in the text. John 
Thieret eventually became an editor for Flora of North America, and the edito- 
rial board adopted the term pistil over carpel. 1am sure it was difficult for him 
to accept such a change, but he did. As it turned out, he was the taxon editor for 
the families that I contributed to FNA. He had to fight with me to get me to use 
the term pistil. I told him why, he agreed, but he said his hands were tied. I told 
him I was not going to type in the word pistil, but he was welcome to replace 
carpel with pistil in my manuscripts, which he did reluctantly. 

One of the courses that Thieret taught at USL was Plant Ecology. I had pre- 
viously taken such a course, so I did not take it from Thieret. He decided to take 
the students on a field trip to Western Texas and Eastern New Mexico. He had 
never been to that area, but I had been there several times, as my in-laws lived 
in Eastern New Mexico at the time. He asked me where to see habitats, and | 
told him several areas that would be good. Rather than attempt a trip on his 
own, he asked me to serve as a guide on the trip. I agreed, and we headed west. 
We were in the desert scrub west of Sanderson, Texas one afternoon where he 


28 BRIT.ORG/SIDA 22(1) 


was talking to the class about Ephedra and other desert scrub plants. | had 
moved away from the class and was examining closely some Ephedra plants, 
when he started the class back toward the car. He was moving at his normal 
pace “full steam ahead and damn the torpedoes” when I heard an awful scream 
from him. I was 25 feet or more behind the class when | caught him running! 
He had walked upon a Western Diamond-back Rattlesnake! It was coiled un- 
der some scrub, and he had almost stepped on it. We stood still (after I settled 
his nerves) and watched the snake for a few minutes. It finally began to move, 
and crawled across our path. That was the longest snake that I have ever seen, 
or so it seemed at the time. 

Dr. Thieret wanted to show the ecology class, in addition to desert scrub, 
mountain vegetation, terminating with tree line and above. He asked if such 
existed in southern New Mexico, and I said I could take him above tree line. He 
thought it was too far south, and, as he often did, would occasionally throw 
some joking jabs into the conversation indicating his disbelief. We started to- 
ward Ruidoso, New Mexico, where | knew there wasa ski resort and the moun- 
tain went above timberline. As we turned a curve and the mountain came into 
view, with the alpine tundra and snow cap, he immediately said, “I told you it 
was here, you just would not believe.” Of course it was a joke, and he used this 
phrase very often. Unfortunately, the ski lift was closed for the season, so we had 
to climb up one of the ski runs to timberline. We did so, and enjoyed collecting in 
the alpine tundra, the first time that | actually ever went above timberline. 

Howard Clark and Billy Dan Hinton graduated my first year at USL. David 
Dike came to USL asa graduate student during my second year, and he chose to 
study the life history of Ottelia alismoides (Hydrocharitaceae), a species intro- 
duced from Southeast Asia. The species was known to occur in The Pool. 
Lacassine National Wildlife Refuge, south of Lake Charles, LA. Dr. Thieret, David, 
and I went to Lacassine to plan his study. We were in a small boat in The Pool 
and were being blown rapidly by the wind. 1 saw something in the water that | 
did not recognize and made a grasp at it as we were blown by. Remarkably, | 
grabbed the plant and enough of it was uprooted to make a couple of speci- 
mens. | looked at the specimen and had no idea what it was. | gave it to Dr. Thieret, 
and he, too, had no idea what it was. We placed it ina bag and took it back to the 
lab, where, using Aquatic Plants (of India), I quickly determined it as Blyxa 
aubertii. Hydrocharitaceae). We had gone to study one introduced aquatic from 
Southeast Asia and had found a second. We prepared a manuscript to report 
the species new to North America, which appeared in SIDA later that year. Dr. 
Thieret sent a duplicate to MICH asa gift, and Ed Voss sent back a letter stating 
that was the first record from the Western Hemisphere of Blyxa in their 
herbarium. Thieret immediately sent back a letter indicating that it was our 
first record also (always a practical joker!). 

John Thieret mastered a technique for rapidly pressing specimens and re- 


a 


HAYNES, JOHN W. THIERET, A STUDENT'S PERSPECTIVE 29 


cording the data. While he was working on a Flora of Louisiana, he was collect- 
ing abundantly all over the State. He had us graduate students (usually me) 
prepare his pressing papers. All papers were separated and carefully folded. Any 
that had color printing must be discarded as the color could get on the plants 
and affect the corolla color. He had the carpentry shop on campus prepare him 
a box with two compartments, each compartment slightly larger than the size 
of a folded sheet of newspaper. This box was a permanent fixture in the rear of 
the camper mentioned above. 

We graduate students were to divide the folded newspaper into two stacks, 
one on which we stamped his collection numbers and the other without num- 
bers. We used the herbarium numbering machine to stamp his collection num- 
bers on the paper. The numbered stack went into one side of the box, and the 
unnumbered stack went into the other side. In the field we would collect a batch 
of specimens and stack them beside the box. 

Thieret always pressed his collections within a few minutes following col- 
lecting. After a short time following collection, he would say, “we have a press- 
ing engagement.” That meant it was time to press plants. He hada tape recorder 
that he used to record the data, and this tape recorder always rode in the box, 
with the newspaper serving as padding for it. He would stand in front of the 
box of papers, record the collecting locality and habitat data, get a numbered 
paper from top of the stack, read that number into the recorder, record the spe- 
cies name, and finally record any specific information about the specimen. The 
first individual was pressed in the numbered paper, and all duplicate speci- 
mens were pressed in unnumbered papers. It was always this sequence (consis- 
tency!), so keeping duplicates with the correct number even though they were 
in unnumbered papers was easy. The field pressing was in a field press, com- 
posed of six or seven cardboards cinched with a press strap. The full field press 
stayed in the truck the remainder of the trip, and we transferred specimens toa 
permanent press upon returning to the lab, often four or five days later. 

For the evening, we would park wherever he could find acceptable and 
spend the night. This place may be a roadside park (illegal in Louisiana, but 
with State plates he always got by with it), state park, etc. One of his favorite 
places to spend the night was a rural cemetery. He said that no one visited these 
cemeteries at night, and he would not be bothered. I do not think he ever was. 
Each night in the field, all of the recorded data were transcribed. He had a por- 
table typewriter that stayed in the camper, and he would type all data of that 
day into his loose-leaf field notebook. The recorder then was ready for the next 
day. Once back at the lab, he had a secretary type labels using his typed field 
notes. All duplicate labels were photocopied, a practice that probably was not 
advisable with technology of the day. Also once back at the lab the process of 
moving specimens from field press to permanent press began. Again, consis- 
tency was important here! One would always begin at the TOP of the field press, 


30 BRIT.ORG/SIDA 22(1) 


never the bottom. (Remember here the numbered paper went into the field press 
first, followed by unnumbered papers.) Starting from the top, then the unnum- 
bered papers would go into the press first for a particular collection, with the 
final sheet of a collection being numbered. Consequently it went on top for that 
collection number. After all specimens were transferred to permanent presses, 
then the presses were placed on the driers, which were in the work area of the 
greenhouse. Removing the specimens from the permanent press also required 
consistency. Always start from the TOP never the bottom. As the press was dis- 
assembled, the first paper out was numbered, usually followed by one or more 
unnumbered papers. These unnumbered papers went inside of the numbered 
one until another numbered paper arrived. This process was repeated until all 
specimens were out of the press. Using this procedure, regardless if the same 
species immediately followed itself, the duplicates always stayed together. Af- 
ter the presses were broken down, the specimens were all placed in a cabinet for 
unmounted specimens. 

Collections he made for the Flora of Louisiana project were used as ex- 
change specimens for the USL herbarium (LAF). Once the unmounted cabinet 
was full or nearly so, which was once or twice a year, he and I would begin the 
process of deciding where to send duplicates. He exchanged with 15 or 20 her- 
baria. We would spread out the numbered paper followed by all of the unnum- 
bered ones of a collection. The best specimen would go into the LAF stack, and 
then all others were sorted into one, two, three, four, etc. stacks, one sheet per 
number per stack, one stack per exchange herbarium. The original label went 
in the sheet for LAF, and photocopies went with the duplicates. It was my re- 
sponsibility to get the duplicates all boxed, shipping labels and invoices pre- 
pared, and get the specimens to the post office. 

With us sending out thousands of exchange specimens each year, we also 
received thousands of exchange specimens in return. Thieret and I often made 
a game of opening the exchange boxes. We would turn the stack so that the 
labels were face down and both stand there in anticipation. One of us would 
quickly open the top newspaper, and our game was to see which one could say 
the name first. He usually won, but I did occasionally - he probably just let me 
win, however. 

After I graduated from USL, l attended The Ohio State University, where I 
earned my Ph. D. under the direction of Dr. Ronald L. Stuckey. While I was at 
Columbus, Dr. Thieret was hired by Northern Kentucky University, in Alexan- 
dria, KY to be Chairman of Biological Sciences. 

John Thieret believed in precision. He always said he was easily pleased 
with perfection. Once I had completed my Ph.D., Dr. Thieret invited me down 
to give a seminar at NKU. Since he had known my wife and son while we were 
in Lafayette, he invited us to stay at his home in Alexandria. He sent directions 
that stated after crossing the Ohio River in Cincinnati continue south until in- 


HAYNES, JOHN W. THIERET, A STUDENT’S PERSPECTIVE 31 


tersect State Route 27. Then follow State Route 27 to Alexandria. After we crossed 
the Ohio River, we noticed U. S. 27 a mile or so from the bridge. | told my wife 
that Dr. Thieret was so precise that he certainly did not mean U.S. 27 since he 
clearly stated State Route 27 (we had his directions in handl!). We continued 
south and finally gave up, deciding that he must have meant U.S. 27, which he 
did. We turned around, found U. S. 27 and followed his directions onto campus. 
He was outside the Biology Building frantically awaiting our arrival, since it 
was less than 30 minutes before the seminar was scheduled to begin. His his- 
tory of precision almost upset the applecart. 

I saw Dr. Thieret only occasionally after I left Lafayette, although we al- 
ways kept in touch. Following the start of FNA, where I ran across him mostly 
was at the Missouri Botanical Gardens, as he was always there just prior to the 
annual Systematics Symposium for a FNA editorial meeting. In July 1998, he 
and I made a two week collecting trip to Utah and Nevada. We agreed that I 
would drive my Jeep SUV and that I would pick him up at the St. Louis airport. 
I was to bring all of the collecting equipment and floras since | had much more 
room than he would have. When I met him at the airport, I was surprised to see 
that he had one small shoulder bag for a two week trip. | asked about his lug- 
gage. He said this is all I have. Needless to say, I could not image how he was 
going to get along for two weeks with just that in the small bag (certainly no 
larger than 3 inches by 9 inches by 15 inches). He said it was less expensive for 
him to fly from Lexington to St. Louis than from Cincinnati to St. Louis, regard- 
less that the plane leaving Lexington stopped in Cincinnati prior to arriving at 
St. Louis and similarly on return. So Mrs. Thieret drove him to Lexington where 
he caught the plane and flew to St. Louis. His plans were to exit the plane in 
Cincinnati on the way home rather than go to Lexington. He obviously could 
not get his luggage if it was checked to Lexington, so he had to take aboard 
everything he needed. I stillam amused about his reasoning, although it worked. 
We had a wonderful time in the mountains and deserts. | collected many aquat- 
ics, and he collected many grasses (all under my numbers). I bet my friends 
who know | almost never collect grasses could not imagine why so many ap- 
peared under my numbers! Regretfully, the last time I ever saw him was when | 
left him at the airport in St. Louis after that wonderful two weeks. | did talk 
with him occasionally, and we did often exchange e-mails, but | was never in 
his presence again. 

[regret that we were unable to get together for eight years prior to his death. 
We had planned another collecting trip, this one to Northern Minnesota. My 
mother, however, became seriously ill, and we had to cancel that excursion. 

Few people have affected my life the way John W. Thieret did. We certainly 
had some rocky times when I was handling the herbarium at USL, but we had 
many, many wonderful times together, which completely obliterate any rocky 
times we might have had. 


JOHN W. THIERET, 
VALUABLE BOTANICAL FRIEND (1965-2005) 


Ronald L. Stuckey 


Emeritus Professor of Botany 
Herbarium, Museum of Biological Diversity 
Ohio State University 
1315 Kinnear Road 
Columbus, Ohio 43212-1192, U.S.A. 


John W. Thieret was one of my valuable botanical friends, and his loss through 
death is now clearly becoming a reality. Il received notice of his death on 7 De- 
cember 2005 from a short morning telephone conversation from his zoological 
colleague, Tom Rambo of Northern Kentucky University. Tom has been known 
to me since his graduate school days when he was studying ornithology. We 
were together during summers at The Ohio State University’s Franz Theodore 
Stone Laboratory Biological Field Station in western Lake Erie on Gibraltar and 
South Bass Islands, at the town of Put-in-Bay, Ohio. My recollections of John 
Thieret are based on facts that I have obtained from my correspondence with 
him, students of mine that have taken his Field Botany class that he taught at 
the Stone Laboratory, my interactions with him as an editor for Sida and book 
review editor for Economic Botany, and other recollections I have of him during 
our 40 years of working together. 

In 1965, | had completed my doctoral dissertation on The Taxonomy and 
Distribution of the Genus Rorippa (Cruciferae) North America, a group of yel- 
low-petaled plants in the mustard family, commonly referred toas marsh cress. 
I received the Ph. D. degree at graduation on 1 May 1965 at the University of 
Michigan, Ann Arbor, and arrived in Columbus in September of that year to 
begin my career as an assistant professor of botany at The Ohio State Univer- 
sity. | believe I had heard of the name John W Thieret, as a botanist who had 
been in the Chicago area, but now was teaching and conducting research at the 
University of Southwestern Louisiana at Lafayette. 


John W. Thieret, Rorippa, and Ronald L. Stuckey (1965-1966) 

A letter dated 3 December 1965 arrived addressed to me, the short text which 
stated: 

We note with interest that you are carrying on work with Rorippa. May we send to you our speci- 
mens of this taxon for annotation? Most of the material in our herbarium consists of Louisiana plants 
collected during extensive field work over the past several years in connection with our state flora 


project 


SIDA 22(1): 33 — 49. 2006 


34 BRIT.ORG/SIDA 22(1) 


I responded on 7 December (exactly 40 years earlier on the same day of his death) 
telling him that I would annotate his specimens of Rorippa, and on 28 January 
1966 | signed the loan form for 31 specimens and one to retain. Upon returning 
the loaned specimens, I wrote in my letter to him of 1 March 1966 offering to 
write a key to the species of Rorippa that grow in Louisiana, as John was pre- 
paring a flora of that state. | also noted that my dissertation was not yet pub- 
lished, and therefore some new names and/or combinations were not yet valid. 
John’s immediate reply of thanks for my annotations on the specimens was re- 
ceived a few days later. The letter concluded with “Let us know if we can ever be 
of help to you. We are always willing to loan or to try to collect specimens for 
those who ask...” 


John W. Thieret and Robert R. Haynes (1968-1969) 

Robert R. Haynes was a graduate student of who worked in the Herbarium un- 
der John Thieret’s direction. Bob also earned a masters degree at the University 
of Southwestern Louisiana, Lafayette with a thesis, titled “A Monograph of the 
Genus Conopholis (Orobanchaceae),” completed in May 1969. Robert was re- 
ally more interested in aquatic vascular plants, and he wanted to study pond- 
weeds in the genus Potamogeton but, of course, no Ph. D. program was offered in 
biology or botany at the University of Southwestern Louisiana. He sought 
Thieret’s advice, who told him that a young professor by the name of Ronald L. 
Stuckey in the Department of Botany at The Ohio State University might be a 
place where he could study these narrow-leaved pondweeds. 


Robert R. Haynes, Potamogeton, and Ronald L. Stuckey (1968-1973) 
Bob’s first letter dated 12 October 1968, came to me with the following request: 


A f 


This spring I hope to receive a Master of Sci the University of Southwestern Louisi- 
ana.I plan to begin work this summer toward a Detar of Philosophy degree in plant systematics and 


am considering Ohio State University as a possible school. My main interests lie in aquatic plants. 


= 


Pret — | would like to do some biosystematic work in the genus Potamogeton subsection Pusilli 
m writing to inquire if there is any possibility of working under your direction and of oe 
oe the financial aid that will be necessary if lam to continue my studies 


By this time Bob had in press his paper on the “Potamogeton in Louisiana” (Proc. 
Louisiana Acad. Sci. 31: 82-90. 1968). The first paragraph of my reply to Bob on 
22 October 1968 stated: 

Itisa pleasure to learn of your interest in beginning a program toward a Doctor 
of Philosophy degree in plant systematics at The Ohio State University. | am 
also pleased to learn of your interests in aquatic plants, particularly in the ge- 
nus Potamogeton. 

My letter continued explaining that | was “primarily interested in floristic and 
phytogeographical problems in aquatic and shore plants.” | acknowledged that 
his interest in aquatic plants would fit into our program. I mentioned the avail- 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 35 


ability of teaching assistantships, and asked Bob that he send a summary of his 
course work, his graduate Record Examination Scores, and asked the Graduate 
School Admissions Office to send him application forms and instructions. John 
W. Thieret’s letter of recommendation for Bob’s graduate work has not been lo- 
cated and probably no longer is extant. Bob's application for Graduate School, 
after review by the Graduate Committee in the Department of Botany, initially 
was not favorable for accepting him. John W. Schmit, chairman of the committee 
and I, with the permission of the Committee, agreed to give Bob a trial period, 
and asked him to enroll in the 10-week summer field program, taking a maxi- 
mum of four courses at the Franz Theodore Stone Laboratory. Bob completed this 
work during the summer of 1969 and earned Ass in all four courses. He then re- 
turned to Louisiana where he was a teacher at Cecila Jr High School in Cecila. 

In the fall of 1969, Department Chairman Schmitt and College Dean Rich- 
ard H. Behning agreed that I needed more permanent help with my responsi- 
bilities as Curator of The Ohio State University Herbarium. The position of 
Assistant Curator was established. The first person appointed to the position 
had to leave at the conclusion of the fall 1969 quarter, and with the position 
vacant, I called Robert R. Haynes to see if he would accept that responsibility. 
From February 1970 to June 1971 he was the Assistant Curator in the OSU Her- 
barium, and from September 1971 to June 1973 he held teaching assistantships. 
Bob completed his dissertation in June 1973 on “A Revision of North American 
Potamogeton Subsection Pusilli (Potamogetonaceae),” which was published in 
Rhodora 76: 564-649. 1975. Bob remained in Columbusat The Ohio State Univer- 
sity for the academic year 1973-1974 and was an instructor in botany teaching 
General Botany for three quarters and Local Flora during the summer quarter. 

Bob developed a career as a professor teaching plant taxonomy, advising 
graduate students, curating a herbarium—all of which concluded with his re- 
tirement in May 2005 from the University of Alabama, Tuscaloosa. His major 
work on Potamogeton and related genera is published in the Flora of North 
America North of Mexico, Volume 22: 3-94. 2000. 


John W. Thieret and the Stone Laboratory (1970-1971) 

In 1956 a new summer teaching program was developed at the FT. Stone Labo- 
ratory under newly appointed Director Dr. Loren S. Putnam, ornithologist of 
The Ohio State University. In the new system, Dr. T. Richard Fisher of the De- 
partment of Botany taught two 5-week courses, Field Botany during the first 
term and Higher Aquatic Plants during the second term. I had the privilege of 
attending the Stone Laboratory and took the Field Botany course from him in 
the summer of 1959. Since that time I kept in contact with Dr. Fisher, who later 
was responsible for my joining the faculty of The Ohio State University in the 
fall of 1965. Among my responsibilities eventually was to teach at the Stone 
Laboratory, and that opportunity surprisingly came the following summer 


36 BRIT.ORG/SIDA 22(1) 


when I taught Field Botany. During the 1967 season I taught the Aquatic Plants 
course. Then in 1968 Fisher left the University to become chairman of the Bio- 
logical Sciences Department at Bowling Green State University in Ohio. Dr. Put- 
nam, who hada budget for the courses at the Laboratory, and in agreement with 
chairman Schmit of the Department of Botany, chose to hire a non-OSU teacher 
for the Field Botany course in 1969 and 1970. That arrangement was to allow 
me to have more time to develop a research program studying the aquatic vas- 
cular plants of western Lake Erie, instead of teaching both courses as Dr. Fisher 
had done since 1956. Putnam also preferred to have non-OSU professors teach- 
ing two consecutive summers and then change to a different teacher. Putnam 
relied on my advice in these situations and asked whom we might hire to teach 
the Field Botany course during the summer of 1971. At the time Robert Haynes 
was the Assistant Curator of the OSU Herbarium, and | asked him for sugges- 
tions. Naturally he suggested John W. Thieret, and I reported to Putnam with 
this recommendation 

From Putnam | soon knew that Thieret was hired to teach Field Botany, 
and a letter of 9 November 1970 came to me from Thieret: 


Ive just spoken with Dr. eee epenaine Put-in-Bay next summer. Lasked him to suggest someone 


to whom I could writ rsyll abus or what-have e-you u for Botany 
610, Field Botany; the text or texts ‘ised: class ne pies ocalities; etc.; etc.; etc. In response 


to my question, he suggested your name. Thus you are the chosen victim. 


John followed with four questions that concerned the course outline, the text- 
book used, class procedure, and good field localities to see and collect plants. In 
regard to the latter item he wrote: 
| must admit that lam a bit panicked by the thought of trying to find enough such localities on a 
one island. (A bit more limited area than lam used to at Lake Itacsa, you know. | mean “Itasca.”).. 

n certainly looking forward to trying my hand at Put-in-Bay. Maybe I'll even be given the shandees 
meet you. Best wishes. And please give my regards to Bob Haynes. 


My three and one-half page letter of 17 November 1970 followed. It began: 

lam very pleased to learn that you are going to be with us teaching Botany 610 (Field Botany), this 
summer at Stone Laboratory. Your letter of 9 November, full of questions is at hand, and I'll try to 
answer them as best I can. I took the field botany course at Stone Lab in 1959 and taught it once, in 
1966. Since then I have been developing and teaching the aquatic plant course which meets every 
other summer and to develop a research program around the theme of the changing flora of the 
islands and of the aquatic situations that we find in the area. 


My letter continued by giving John some statements about the kinds of students 
that take the course and their knowledge of plants. I noted that the course prima- 
rily concerned the flowering plants, that Clara G. Weishaupt'’s Vascular Plants of 
Ohio was the book used, and that I had an outline of the course with various 
procedures that I would mail him. Then I wrote a long statement about class field 


— 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 37 


trips, assuring him that he would have plenty of plants and ample places to take 
students. With reference to field work, ladded a statement about taking field trips 
by boat to the various islands in Lake Erie and to the Ohio mainland. 

My letter concluded with comments on the available library resources, labo- 
ratory supplies for the course, materials for pressing and drying collected plants, 
and the information sheets, more commonly known then as “handouts,” that I 
used in the 1966 Field Botany course. The latter I offered to mail to him. I con- 
cluded the letter about my own projected plans for the up-coming summer. 


I will be at the Laboratory all summer myself. ie Hise term neal be for research, at which time I will 


be spending most oot myt time writing. Hopefully, he “Flora of the Erie 
Islands,” and | the “Origin, Changes, and Geo- 
graphical Relationships of the Acuane and Matsh Flora of Nestea a Erie and Northwestern 

Ohio.” During the second term I will k | lam looking forward very 


h to your coming to Stone Lab next summer. Please do not hesitate to write if you have further 
questions. I hope the above comments will be helpful to you. 


John did not hesitate to send a return letter, dated 20 November 1970. It began, 
“I certainly thank you for your long letter—and all the useful data therein. Be- 
lieve me, your help is deeply appreciated.” After commenting on the various 
items | had mailed him, he concluded his letter saying, “After I've gone over all 
the material thoroughly, I'll let you know if any other questions arise.... Thanks 
a lot for all your fine help.” 

John continued his pursuit concerning the 1971 summer program at the 
Stone Laboratory, and on 19 January 1971, he wrote to Director Putnam, greet- 
ing him with: 

Dear Puttie: 

I note in your bulletin for summer 1970 that you had, in 1969 a series of talks—a “Special Lecture 

ae 25 Ca ee ay am not “outstanding” (except in my unrivalled 
lam noscientist, I still intend, via this letter, to offer 


to give some sort of a talk this coming summer. I have done quite a bit of botanical field work in 
Canada’s District of Mackenzie—in the Great Slave Lake region. Asa result of the work, I have a set of 
slides that some people find interesting, especially when the slides are accompanied by the delight- 
ful eee presented by humble John. If _ wish such a presentation—let’s call it a travelog— 
I'd be glad to import my slides to Put-in-Bay this coming June. I'm not at all certain that a travelog by 
as ee be worth a mention in your 1972 eae of it might fill in an otherwise empty evening 
on Gibraltar. And besides, I like to talk..Let me know. 


This letter reveals much of the kind of humor John displayed about himself. 
Regarding these special lectures, beginning in the summer of 1970, Director 
Putnam asked me to make arrangements for the special lecture series at the 
Laboratory. | contacted the potential speakers, prepared the program, hosted 
their visit, and introduced them when they made their presentations on Thurs- 
day evenings. Scheduling a special lecture by John W. Theiret was easily ac- 
complished, and I distributed the lecture schedule with Thieret listed for 13 


38 BRIT.ORG/SIDA 22(1) 


July 1971 with the title “The Yellowknife Highway Region, Canada: Flora and 
Vegetation.” John’s wish to present a lecture was fulfilled, and his lecture title 
later was listed in the 1972 admissions bulletin. 


Again, Thieret, Rorippa, and Stuckey (1966-1972) 

When I completed my dissertation on the genus Rorippa in 1965, it was diffi- 
cult to locate a journal that would publish a large taxonomic monograph as a 
single paper or issue of a journal. My advisor, Edward G. Voss, had anticipated 
that I could retain the entire dissertation as one unit if it were published in the 
Contributions from the University of Michigan Herbarium. This publication had 
been inactive for some years, and editor Rogers Mc Vaugh reaffirmed that funds 
were not available at that time to reactivate it. The conclusion was that 1 would 
have to look elsewhere to publish my dissertation. 

Edward G. Voss held in high regard the taxonomic knowledge of Lloyd H. 
Shinners of Southern Methodist University. When Lloyd taught the field tax- 
onomy course in the summer of 1952, at the University of Michigan Biological 
Station, Pellston, Ed was his graduate teaching assistant and from then on grew 
a close botanical relationship and great respect for each other. In 1962, at SMU 
Shinners began publishing a private taxonomic journal named Sida, and Ed 
suggested that | might contact Dr. Shinners regarding publication of my Rorippa 
monograph. I then proceeded to correspond with Shinners about my study of 
Rorippa, and during 1966 I extracted and published two short papers taken from 
my dissertation. They were “The distribution of Rorippa sylvestris (Cruciferae) 
in North America” (Sida 2:361-376. 1966), and Rorippa walteri and R. obtusa 
synonyms of R. teres (Cruciferae)” (Sida 2:409-418. 1966). During 1967 through 
1969, I revised my monograph on Rorippa, and on 5 December 1969, I wrote to 
Dr. Shinners asking if he would publish my Rorippa study in Sida. He enthusi- 
astically responded on 9 December 1969, and told me to send the manuscript to 
him after Christmas. A letter of 26 March 1970 confirmed that he had accepted 
the manuscript, whose “bulk is a bit appalling.” | heard nothing further from 
Shinners until I learned of his death of 16 February 1971. The future status of 
Sida was in doubt, and therefore the publication of my Rorippa monograph 
also was in doubt. Later on 15 March 19711 wrote, I wrote Dr. William FE Mahler, 
who was Shinners replacement, thanking him for his letter of 3 March 197] 
which indicated that publication of Sida would be continued and that addi- 
tional work was to proceed on my Rorippa monograph. “I am pleased that you 
have selected Dr. Thieret to go over my manuscript in preparation for its 
publication...As you may know Dr. Thieret will be at our University for 5 weeks 
this summer teaching at Stone Laboratory. | will be there at the same time and 
can work with him on the manuscript.” 

In April 1971, John and I began a correspondence on the preparation of the 
Rorippa monograph for publication in Sida. Together, John and I discussed the 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 39 


Rorippa manuscript while both of us were at the Stone Laboratory. Later in the 
year John wrote me letters on the progress of the editing which continued into 
1972. John’s letter of 11 January 1972 stated: 

When I finish with the ms., I shall return it to Mahler. I have just been asked to serve as Associate 
Editor of Sida—which is just what I’ve been doing all along! 


By April John had sent the edited Rorippa manuscript to Mahler, and the latter 
replied to me, 13 April 1972, that my “manuscript is already at the printers.” 
Thieret received the galley proof sheets and then sent them on to me for read- 
ing. Together that summer at Stone Laboratory, John and I read the galley sheets. 
Page proofs came on 25 July 1972, according to Mahler's letter to me, on 8 Au- 
gust 1972. lreturned the final page proofs, and by 27 September, | was complet- 
ing the transaction to purchase 200 copies of my study of Rorippa. My mono- 
graph, “The Taxonomy and Distribution of the genus Rorippa (Cruciferae) in 
North America” appeared in Sida 4(4):279-430. 1972. John W. Thieret was listed 
as the associate editor and William F. Mahler as the copyright owner. Had it not 
been for Thieret entering the situation, my monograph might never have been 
published. I owed hima great debt of gratitude for his unselfish help in editing 
my publication. 


Again, Thieret and Stone Laboratory (1972) 
In the summer of 1972, Thieret returned to the Stone Laboratory and again 
taught Field Botany. This summer he had prepared a book of mimeographed 
course handouts that were to be given to each student. Nearly everything that 
John wrote had some humor associated with it. At the bottom of the Table of 
Contents page was the notation, HAVE YOU THANKED A GREEN PLANT TODAY? 
y masters student David L. Moore took the course from Thieret that sum- 

mer. At that time also, David was studying the changes, since 1895, in the aquatic 
vascular plant-flora of East Harbor State Park, Ottawa County, Ohio (Completed 
1973; portion published in Ohio J. Sci. 76:78-86. 1976). David later in 1976 earned 
the Ph.D. degree from The Ohio State University by studying the distribution 
of freshwater algae in northeastern Ohio with Dr. Clarence E. Taft. Since then 
David has taught botany courses in the Department of Biology at Utica College 
of Syracuse University, Utica, New York. Beginning in 1993, he returned to the 
Stone Laboratory and has taught the aquatic plants course every summer ex- 
cept 2005. The course was not taught that summer. 

In response to my request for information about John Thieret’s 1972 Field 
Botany class, David provided the following three paragraphs: 


As | retrieved an OSU soft cover binder from my office shelf, | was reminded of the passage of 34 


years by the yellowed, brown- edged, | hand-written pa ages! ies summer 1972 Field B otany Course W ith 
Dr. Joka ae came pac tome as | iad SS the pages of notes. We learned a total of 19] taxa 
in those five 25 plant families. Indeed it was the first time I had ever 


seen floral formulae and floral ae My favorite plant was Matricaria matricarioides, the Pine- 


40 BRIT.ORG/SIDA 22(1) 


apple Weed, which Dr. Thieret picked by the docks and gleefully crushed in his palm so we could 
sniff the fragrance of fresh pineapple. 


At the first class session Dr. Thieret went over the syllabus. Everyone laughed 
when he referred to examinations as “mental enemas.” I will always remember 
the cover of his mimeographed course handouts which he provided to us that 
first day. The front page pictured Shiva Nataraja, the Hidu Lord of the Dance 
ona Nuphar advena leaf which protruded from the fruit of Nelumbo lutea. In the 
right rear hand was a leaf of Rhus toxicodendron (Poison Ivy) while in her left 
rear hand was an Aescylus glabra leaf (Buckeye). The afternoon of July 8 was 
my first trip to Pelee Island. It was probably one of the highlights of the session, 
and where I first saw Populus grandidentata and Diplotaxis. | still don’t know if 
it was D. tenuifolia or D. muralis. 

At the end of the session, just before the final exam, Dr. Thieret read a poem 
he had composed on the porch of Bayview House where he and his family lived. 
In it he dedicates a verse to each of the students in the class. After he read it he 
gave each of us a two page mimeographed copy on yellow paper. I kept my copy 
in my notebook after we finished the course. | retyped it with the same spelling, 
punctuation and format that he used except that I placed it in two columns. 
There is one misspelling left intact - “cannister”- and the two lower case letters 
which should have been capitalized. All of his “poetic license was retained). 

| have selected the following six verses of John’s poem, titled “Botany 610 In 
RHYM, Summer 1972,” for publication here: 

BoTANy 610 IN RuHyYM 
SUMMER 1972 


This is the forest primeval That lovely young Madchen Miss Schiller 


The murmuring Acer and Celtis 
is is the class that beneath it 


Romped cheerfully after their Leader. 


This is the class below pictured 


In verses rhythmic, poetic. 


‘Twas a pleasure to watch Mr. Moore 


Whose knowledge of plants was not poor. 


Like an eye-talian barber 
He combs over east Harbor 
And there gave Fearless Leader a tour. 


And now it is time for Miss Kneller 
[tried hard this plant world to sell ‘er. 
By selling my wares 

lincreased her cares. 

So often “Work hard, dear” Id tell ‘er. 


With plant names | really did fill ‘er. 
But a turtle so cool 

She found ina pool 

Was really, | think, what did thrill ‘er. 


There goes that bold man Rick McGill 
Out 
He works day and night 
Wit 
And does get his plant press quite fill. 


ora botanical kill. 


vallof his might 


And last, but not least, Fearless Leader. 
Whose disciples learned oak, elm, and cedar. 
How oft, on The Rock 

He did gather his flock 

And exhort them there, much like old Peter. 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 41 


At the conclusion of the poem, John 
wrote, “... presented with thanks and 
appreciation to my Stone Lab Students. 
Long may they flower!” 

The verse pertaining to Rick McGill 
is noteworthy as he was working with 
me for his master’s degree making acom- 
parison of the vascular plant flora of the 
two lakes in northern Champaign 
County, Ohio (completed 1973; not pub- 
lished). 


Field Botany, 
Course Handouts 


Summer 1993, 


John W. Thieret, Tom Duncan, and the 
“Flora of the Erie Islands” (1972-2005) 
Tom Duncan, an undergraduate major- 
ing in education and working asa teach- 
ing assistant in chemistry at the The 
Ohio State University, introduced him- 
self to me during his sophomore year in 
the early spring of 1968. Later that spring 
he accompanied me on field trips learning the local flora, and in the spring of 
1969 he began working as an assistant under my direction in the OSU Her- 
barium. That summer I invited Tom to serve as my research assistant at the FT. 
Stone Laboratory. He continued to serve part-time as my research assistant on 
the Flora of the Erie Islands until he graduated at the end of the fall quarter 
1970. Tom continued working on the Flora manuscript from 1971 through 1973 
while he was a graduate student at the University of Mighican. He earned his 
Ph. D.in Botany there in 1976 and was employed as an assistant professor botany 
beginning 1 July 1976 at the University of California, Berkeley. He was granted 
tenure there in 1982, was the Director of the Herbarium 1982-1991, then be- 
came Director of the Museum Informatics Project until his retirement in 2002 
as an associate professor emeritus at age 54. 

I first met John W. Thieret in July 1972 at the Franz Theodore Stone Labo- 
ratory on Gibraltar Island in Lake Erie. John was at Stone Laboratory for the 
summer to teach Field Botany. I was visiting Stone Laboratory for a few days to 
discuss with Ronald L. Stuckey progress on the preparation of our manuscript 
titled The Flora of the Erie Islands: Its Origin, History, and Change. 

Stuckey initiated a project on the flora of the Erie Islands in the mid-60s. In 
1969, I undertook a project as his research assistant on the changes in the flora 
of the seven small islands in western Lake Erie (The Michigan Botanist 9:175- 
200. 1970), and was invited to collaborate on the more inclusive project in 1970. 


Cover page of John W. Thieret’s Field Botany, Course 
Handouts, Summer 1972. 


42 BRIT.ORG/SIDA 22(1) 


By 1972 we were in the initial stages of preparing together a manuscript de- 
scribing the results of our research. 

John Thieret was spending his second summer as an instructor at Stone 
Laboratory. He had become intensely focused on the flora of the islands and 
was interested in discussing with me the progress to date on my collaboration 
with Stuckey in this regard. One area we discussed was the lack of information 
in previous floras about cultivated plants on the islands. 

Cultivated species are of interest because they are an important source for 
the adventive and naturalized flora of the islands, are a conspicuous aspect of 
the flora, and are an important feature of the flora for the residents of the is- 
lands. John and | concluded that we would undertake a survey of gardens on 
South Bass Island and compile a list of the cultivated flora for inclusion in the 
manuscript Stuckey and | were preparing on the total flora. 

We spent a day wandering the roads of South Bass Island stopping at houses 
and asking local residents if we could examine and record the names of the 
plants in their gardens. With a copy of Liberty Hyde Bailey’s Manual of Culti- 
vated Plants (1949), in hand, we were able to prepare a preliminary list of plants. 
Ultimately this list included 258 taxa. 

John often stated that our field trip on South Bass Island that day was one 
of his favorite memories of his time at Lake Erie. After our initial meeting in 
1972 we continued to correspond and talk on the telephone about the progress 
of the flora manuscript. In 1982 John brought the nomenclature of our list of 
cultivated plants up to date using Hortus Third (1976), the nomenclatural ref- 
erence for cultivated plants at that time. In his letter of 3 February 1982 he con- 
cluded by saying, “best wishes from John. I wish we could wander over South 
Bass Island again. Fun, fun, fun.” 

Whenever | think of these events, lam always reminded of John’s charm- 
ing personality, his appreciation of natural history, and his joy in studying plants. 
His interest in the flora of the Erie Islands continued for over 30 years until the 
end of his life. 

Stuckey and I completed our manuscript in 1976 and submitted it to the 
Ohio Biological Survey for publication. The Survey accepted the manuscript 
for publication and planned for publication in 1979. John volunteered to serve 
as editor for the Survey and worked extensively on the manuscript. Unfortu- 
nately, the Ohio Biological Survey could not publish the manuscript at that time, 
partially because of the lack of funds, and Stuckey’s involvement with other 
projects for the Survey. For the next 20 years the manuscript and three edited 
copies resided in Stuckey’s office with an additional copy in the Franz Theodore 
Stone Laboratory library. 

During this period, John continued his interest and frequently asked both 
Stuckey and me if we would complete the manuscript and find a way to render 
itin a form for wide distribution. During a visit to The Ohio State University in 


— 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 43 


1999, Stuckey and I discussed what might be done to resurrect the manuscript, 
complete it with all editorial suggestions incorporated, and either publish it or 
distribute it electronically. 

John was enthusiastic about this turn of events and offered his services to 
help us in any way he could. Progress in this direction was made but no definite 
arrangements about a means of distribution had been made by the time of John’s 
death. We are investigating the possibility of distributing it through the digital 
Knowledge Bank at The Ohio State University. Ronald L. Stuckey and | are de- 
termined tocomplete this task to honor John’s long-standing interest in the Erie 
Islands, and we plan to dedicate the completed flora to him. 


Thieret and Stuckey’s Promotion (1972) 
Later in the year 1972, John made an important contribution to my career. He 
wrote a letter, dated 19 October, supporting my promotion to full professor in 
The Ohio State University. Addressed to Botany Department Chairman John A. 
Schmitt, it was short and to the point, quoted here in its entirety: 
It seems to me that Dr. Stuckey has shown himself to be a competent and indefatigable researcher. 
His frequent publications attest to his capabilities not only as an “original researcher” but asa skilled 
synthesizer of the work of others. 

[PS] John: This is a trite, trivial, and, for me, traumatic letter. | simply cannot rise to greater 
heights. Even though the subject is worthy of those greater heights. Jlohn 


Again, Thieret and Stone Laboratory (1974, 1977, 1981, 1983, 1986) 

After 18 years as Director of the Stone Laboratory, Putnam retired following the 
summer session of 1973, and College Dean Richard H. Bohning appointed Dr. 
Charles E. Herdendorf to succeed him. Now Professor and Director Emeritus, 
Herdendorf, on 23 March 2006, provided his remembrances of John Thieret, 
briefly reviewing the succeeding summers John taught Field Botany at the Labo- 
ratory—a total of seven summers over a 15 year period. 

I first met Dr. John W. Thieret, in 1971 at the Stone Laboratory where he was 
teaching a summer course in Field Botany, and I was beginning to organize the 
newly founded Center for Lake Erie Area Research (CLEAR) at the Laboratory. 
The next year we both served on the summer faculty of Stone Laboratory, Dr. 
Thieret again teaching Field Botany, while I taught Physical Limnology. Finan- 
cial constraints in 1973 caused the cancellation of classes at the Laboratory, but 
by 1974 I had assumed the duties of Laboratory Director, and Dr. Thieret was 
invited again to teach field botany. In 1974, Dr. Thieret and his class also par- 
ticipated in a CLEAR research project titled Ecological Analysis Related to the 
Proposed Seawall Improvement and Ground Rehabilitation Project at Perry’s Vic- 
tory Monument, Put-in-Bay, Ohio. Dr. Thieret returned to Stone Laboratory to 
teach Field Botany in 1977, 1981, 1983, and 1986. On June 30, 1983, he also par- 
ticipated in the Stone Laboratory Guest Lecture Series by presenting a seminar 
titled The Muskegand |: Exploring for Plants in Canada’s Northwest Territories. 


44 BRIT.ORG/SIDA 22(1) 


Other than his account of the boreal plants along the new Yellowknife High- 
way, one of his most memorable descriptions was that of the voracious infesta- 
tion of black flies encountered around Great Slave Lake. John Thieret is fondly 
remembered for his dry sense of humor and fascinating stories of field experi- 
ences. Some of his students may have found him to be excessively demanding, 
but all received a quality introduction to the study of Field Botany. 

I was not present at the Stone Laboratory in the summer of 1977, but my 
masters graduate student John R. Wehrmeister attended and took the Field 
Botany course from Theiret. Student Wehrmeister was researching the ecologi- 
cal life cycle of the pondweed Potamogton crispus in North America (completed 
1978; published in part The Michigan Botanist 31:3-16. 1992). In 1981, Mr. 
Wehrmeister earned an M.D. degree from the Medical College of Ohio in To- 
ledo, and he has had a successful career as a physician in the practice of inter- 
nal medicine in the Maumee-Toledo, Ohio, area. In reply to my request, he wrote 
the following commentary: 
lam pleased to offer thoughts about Dr. Theiret. His personality and teaching style has had a lasting 
1m 


act on me. | had heard Dr. Thieret give a lecture on the botany of the Northwest Territories,Canada 


= 
> 
pane 


was a student at the University of Michigan Biological Station at Pellston. Dr. Theiret was a 
visiting faculty member there in 1975, but I did not get to know him that summer. Later in 1977, 
while studying at The Ohio State University, I took his Field Botany course at the Stone Laboratory. 
Following the course, in late August and early September, my friend Ed Toth, and | accompanied Dr. 
Thieret on a plant collecting trip to Louisiana, and upon returning, visited with him at his home in 
Highland Heights, Kentucky. 


Thoughts of him are full of fond memories. He was a delightful man, a favorite 
but serious teacher, and a scholar with a great sense of humor. Memory of his 
facial expression includes what seemed to be a permanent grin. “Dr. Thieret 
anecdotes” were the order of the day among his students. Good-natured though 
he was, in contrast he was strongly serious in the classroom. With some embar- 
rassment | remember being admonished by him for the poor effort | showed in 
mounting herbarium specimens for an early class project. At my request Dr. 
Theiret wrote a letter of recommendation for some job or school position to 
which I'd applied. He thereupon sent mea purported carbon copy of the letter 
which, perhaps needless to say, was far from complementary. Luckily this was 
one of his pranks. The real letter contained the usual laudatory comments. 

A few additional observations of Theiret’s summers at the Stone Labora- 
tory are noteworthy. By 1981, he had modified the “Course Handouts” book. On 
the cover that summer was a branch showing the leaves of the hackberry tree, 
one of the most common trees that grew on the shallow limestone and dolo- 
mite bedrock. On page 4 was the quotation, “The beginning of wisdom is to call 
things by their right names.—Oriental proverb.” 

In his course the students only were required to learn the scientific names 
of the plants. I did not agree with him; rather I taught students to learn both the 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 45 


John W. Thieret’s Field Botany Class at the F.T. Stone ii etal eke rarer in oe Dare eee quarry 
on Kelley ‘s Island, Erie County, Ohio, by Ronald L. Stuckey, Ju uly hn W. Thieret, George 
F, Buddell, IL, assistant; five men students, in alphabetical ne not matched to individual i in the ie Ron 
Baker, Doug Brewer, John George, Mark Harrell, and Dan Wojta 


scientific and common names, which would allow them to communicate with 
persons of many different educational backgrounds and professions. His edito- 
rial style gave me concern also, but I never discussed it with him. He continu- 
ally overused the dash, which therefore changed my writing style. I also be- 
came annoyed when he was editing the “Flora of the Islands.” | often listed 
several plants in sequence. He would change the sequence by aligning them in 
alphabetical order. My choice of sequence usually was to list them from the 
most common or typical to the less common in a particular habitat. My align- 
ment was ecological related as to where the plants grew. Plants do not grow in 
places alphabetically, but he apparently was not aware that my order of the 
plants also had a meanin 

John liked to talk as he himself has written. During two other summers he 
gave a presentation in the Guest Lecture Series, on 19 July 1977 and 30 June 
1983. Both times, his title was “The Muskeg and I,” which was just as delightful 
as it was the first time with the more formal title, when he gave it in the sum- 
mer of 1971. 

Andrea Wilson, now Mrs. Matthew Schlageter in the Southeastern Devel- 
opment Office of The Ohio State University, Bonita Springs, Florida, came in 


46 BRIT.ORG/SIDA 22(1) 


the summer of 1982 as Director Herdendorl’s Laboratory Secretary and Admin- 
istrative Assistant. She knew John Thieret from the 1983 and 1986 summers at 
the Laboratory. Her statement sent to me 24 March 2006 concisely summa- 
rizes John’s presence there. 

| vividly recall Dr. John Thieret as a very kind and thoughtful man. He al- 
ways had an encouraging word and a smile for everyone. He was devoted to his 
ne touched. 


— 


teaching and research and was well respected by all whose lives 
His memory will continue to live not only in his publications, but also in the 
knowledge he shared with his students. 


Thieret, Book Reviews, and Stuckey (1973-1979) 
During the five year period of 1973-1979, Thieret sent me books to review for 
the journal Economic Botany. For 25 years, 1959-1984, John voluntarily served 
as book editor for Economic Botany. John’s strive for perfection gave the journal 
“a reputation as having one of the most comprehensive and best edited book 
review sections of any professional journal. When at times the acid pen of a 
reviewer might be too harsh on an author, John would always suggest a critical 
but more even-handed approach.” (W. Hardy Eshbaugh, Economic Botany 38(4): 
498. 1984). 

John usually sent me local flora or plant taxonomy books to review. Ini- 


tially he sent me two books with a letter dated 24 September 1973. The book 
titles were A Flora of the White Mountains, California and Nevada by Robert 
M. Lloyd and Richard S. Mitchell 1973) and The Genus Lesquerella (Cruciferae) 
in North America by Reed C. Rollins and Elizabeth A. Shaw (1973). Thieret’s 
suggestions were: 


QQ 


Probably the best thing to do so far as the flora is concerned is simply to produce a “notice” of the 
work. A full-blown “review” is not necessary. If you can get the notice and the review to me in about 3 
months or less, I'd be ecstatic. Allot ner 150- 200 words for the flora, about 300 for the mustard 
opus. Of course, if you fee you really need more—or do not need so many—the final decision is yours. 


| wrote 10 book reviews for Economic Botany. They and all of my other book 
reviews are available in my archives at the Hunt Institute for Botanical Docu- 
mentation, Pittsburgh, Pennsylvania. John wrote his own review about his 25 
years as Book Review editor. He considered it as “one singularly devoid of ex- 
citement.” One colleague, with perhaps a touch of jocularity, called him the “dean 
among U.S. book review editors” (Economic Botany 40:24-26. 1986). 


Thieret, Aquatic Plants, and Stuckey (1982-1983) 

[had forgotten that during the summer of 1983 when John was teaching at the 
Stone Laboratory, he asked me to review the book manuscript he was complet- 
ing for the late Ernest O. Beal on the “Aquatic and Wetland Plants of Kentucky.” 
On 26 August 1983 John wrote: 


| hope you remember that you have gone over both major sections of the book, ie., the dicots (sum- 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 47 


mer 1982) and the monocots (summer 1983). The comments you made on the monocots were (as 
were those on the dicots) gratefully peceveds Most of them have been incorporated into the book, 
| 


improving same! | appreciate your r prose. I’m working hard on the book, hop- 


ing to meet a deadline of | November 1983 ibe getting he book to the publisher. Best wishes. 


I further helped John with the book by writing on 2 September 1983 a support- 
ing letter for its publication to Richard Hannan of the Kentucky Nature Pre- 
serves Commission: 

“Dr. John Thieret has asked me to write in behalf of his manuscript co- 
authored with the late Dr. Beal on the aquatic and wetland plants of Kentucky. 
This manuscript is very carefully prepared in all aspects. Dr. Thieret has gone 
to great lengths to check the accuracy of identifications of specimens seen, the 
reliability of literature records and citations, and the conciseness and correct 
usage of technical terminology. His work in this regard is very well done, and 
adds great credibility to a work of this type. 

Very few states have books on their aquatic plants, and this book will be a 
welcome addition to those books on aquatic plants. Perhaps, more important, 
however, is that this book will be extremely valuable in a state which has no 
“modern” general flora. Consequently, this book will be a valuable contribu- 
tion to that “modern” general flora that needs to be generated. Those needing 
floristic information will at least be able to get current information on aquatic 
and wetland species. I trust that you find this contribution worthy of publica- 
tion, as lanxiously look forward to seeing it in print 

In the book’s acknowledgements by Thieret, my name, along with many 
others, appears among those who “gave aid during the preparation of this book.” 


Thieret and Stuckey Since Summers at Stone Laboratory (1996-2005) 

Since the summer days of the 1970s and 1980s at the Stone Laboratory, most of 
my contacts with John Thieret have been by telephone, except the following 
interactions which are noteworthy. When John became editor of the Transac- 
tions of the Kentucky Academy of Science, [thought he could help me with pub- 
lishing some of my writings that were nearly completed on the controversial 
North American botanist Constantine Samuel Rafinesque (1783-1840). | had 
published several papers on Rafinesque’s botanical studies earlier in my career. 
Since Rafinesque had held the position of Professor of Botany and Natural His- 
tory at Transylvania University in Lexington, Kentucky, from 1819-1826, my 
contributions were appropriate for the Kentucky Academy’s publication. With 
the assistance of my botanical friend, James S. Pringle of the Royal Botanical 
Gardens in Hamilton, Ontario, Canada, we published, with John’s editorial as- 
sistance, “Common names of vascular plants reported by C.S. Rafinesque in an 
1819 descriptive outline of four vegetation regions of Kentucky” Clrans. Ken- 
tucky Acad. Sci. 58:9-19. 1997). At the Bicentennial Celebration of Rafinesque’s 
200 birthday held on the campus of Transylvania University, 21-22 October 


48 BRIT.ORG/SIDA 22(1) 


1983, | spoke on “Opinions of Rafinesque expressed by his American botanical 
contemporaries” (Bartonia 52:26-41. 1986; reprinted with some changes in 
Charles Boewe (Profiles of Rafinesque. Chapter 10. 2003). Following that event, 
and as a contribution toward my studies of the pioneer botanists of the Ohio 
River Valley, | wrote an extensive manuscript on “Rafinesque’s Botanical Pur- 
suits in the Ohio Valley 1818-1826).” With John Theiret’s editorial skills, 14 years 
later this manuscript was completed and published by the Kentucky Academy 
of Science in its newly titled Journal of the Kentucky Academy of Science 
(59(2):111-157. 1998). 

John appreciated and was very supportive of my scholarship on the his- 
tory of nineteenth century North American plant taxonomists and their work. 
He asked me for a short project he could contribute about Rafinesque in Ken- 
tucky. | suggested he return to the localities in the state where Rafinesque ob- 
tained plants that he named as new to science. He was delighted with the idea, 
and by automobile during a week in July 1999, John and his friend, David M. 
Brandenburg of the Dawes Arboretum, Newark, Ohio, retraced a botanical ex- 
cursion that Rafinesque took during 1823 in central and south-central Kentucky, 
Rafinesque’s itinerary was outlined in his autobiography, Life of Travels 1836), 
which took him two months to complete on foot. A short description of the 
Thieret-Brandenburg trip appeared in the Flora of North America Newsletter 
(13:;G-4). 1999. July-December). David and John wrote the full story of the trip 
under the title, “Rafinesque and Us” (Lloydiana 6():4-9. 2001). 

During retirement, my friend George W. Paulson, emeritus professor of 
medicine in the Medical Center of The Ohio State University, has been writing 
about early medical practices of the pioneer physicians in the central Ohio area. 


Among these physicians was Samuel Thomson (1769-1843) who discovered that 


ear 


— 


chemical extracts from the leaves and seeds of the plant, Lobelia inflata, had 
medical properties that potentially would cure headaches, tinnitus, syphilis, 
and especially asan expectorant to loosen bronchial congestion. Paulson wanted 
to write a paper on Thomson, and asked me to help with it since | would know 
about the Lobelia plant. Most of the paper was written by George, and I edited 
it, but George could not locate a journal for its publication, so he asked for my 
help. [thought Lloydiana, a quarterly periodical of articles for the popular au- 
dience published by the Lloyd Library in Cincinnati, would be an ideal place 
for its publication. I talked to John Thieret on the telephone and asked him for 
his review of the article and editorial assistance. John,a member of the Board of 
Trustees of the Lloyd Library, helped with revision and editing of the article, 
which appeared in Lloydiana (8(3):2-6. 2004. Summer). Lloyd Library Direc- 
tor and editor of the periodical noted in her column, “From the Director,” that 
the Thomson-Lobelia paper was indeed an “interesting article.” John Thieret so 
ably helped make it possible to be printed in the Lloyd Library newsletter. 
Nearly every time John and [hada telephone conversation since our teach- 


—_—_ 


STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 49 


ing days at the Stone Laboratory, he always asked about progress on the Flora of 
the Erie Islands, and he wanted me to finish it. | always had to think of some 
excuse, mostly that I was involved in other more worthwhile projects. Some 
relief came for me when Tom Duncan returned to my office in 1999, and he 
agreed to take all of the various components related to the project with him to 
Berkeley, California. There he has worked on the Flora to reflect our knowledge 
of it, as it was in the 1970s. Tom wrote of his role in this effort in this paper. 

For 30 years, from about 1971 to 2001, | was at work preparing another kind 
of book about South Bass and Gibraltar Islands. Titled “Lost Stories,” the book 
includes Theresa Thorndale’s “Island Jottings” about selected places on these 
islands of the 1890s that she published in the Sandusky Register. My contribu- 
tion describes these same places on the islandsa century later during the 1990s. 
This book won the 2003 Florence Roberts Head Book Award given by the 
Ohioana Library Association of Columbus, Ohio. In his last known note to me, 
John Thieret wrote, “Many thanks for the info[rmation] on your award for the 
Put-in-Bay book. ’'m so glad that you did such a book. You certainly deserve the 
award.” I sometimes think that for John, it may have been a consolation prize 
for the Flora of the Erie Islands. 

John Thieret was a most unusual botanist. I know of no one else like him. 
His dedication, his kindness, his sense of humor, his field knowledge of plants, 
and the numerous and varied ways he helped me through my career has been a 
most worthwhile and enjoyable adventure. 


ACKNOWLEDGMENTS AND SOURCES OF INFORMATION 


lam especially grateful and thankful to Kathy Royer for preparing the final 
typewritten manuscript of this paper for publication in Sida. lalso extend grate- 
ful thanks to David lL Moore, Tom Duncan, Charles E. Herdendorf, John 
R.Wehrmeister, and Andrea (Wilson) Schlageter for their written contributions 
to this paper. Notes made during telephone conversations with the above indi- 
viduals and the letters referenced in this paper are filed under the title of this 
publication in the Archives of Ronald L. Stuckey and in the Archives of the 
Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania. As an 
added note, former Stone Laboratory Director Loren S. Putnam died during the 
writing of this paper, 19 April 2006, at age 92.5 years. 


JOHN W. THIERET, 
COLLEAGUE AND EDITOR FRIEND (1969-2005) 


William F. Mahler 


204 W Clara 
lowa Park, Texas 76367-1304, U.S.A. 


John Thieret’s reputation as a dedicated botanist and unique individual pre- 
ceded our actual meeting, which occurred sometime in the late sixties. Over 
the years we drifted apart, but the impact John made on my professional career 
and my success with Sida, Contributions to Botany remains strong. Although 
he spent the night at our house only a couple of times when he visited the SMU 
herbarium, the field trips we took in my 1969 red Ford pickup were the high- 
lights of our botanical relationship. 

Our biggest collecting trip was to Guadalajara, Mexico in April of 1970. We 
traveled by way of the Chihuahuan Desert and then returned through the 
Sonoran Desert. The pickup was the perfect mode of transportation for a long- 
range collecting trip because of the saddle tanks and the camper shell. We could 
travel six hundred miles between fill-ups and could sleep in the back if we 
couldn't find lodging. Also, there was plenty of room for all our collecting sup- 
plies. 

The decision to take this particular trip was made on a lark. We'd heard of 
others doing this and so we thought we'd do the same. As it turned out, we actu- 
ally did find quite a few noteworthy specimens. When Rupert Barneby found 
out where we were going, he suggested we visit some locations for the legumes 
that he had collected some years back. He needed some flowering specimens. 
We were successful in finding some specimens north of Zacatecas, which he 
cited in his monograph on Dalea. 

The trip was far from boring. North of Guadalajara, John injured his ankle 
while trying to get a specimen from the branch of a tree. He knew he had 
sprained his ankle, but he was afraid he might have broken it, too. An x-ray ina 
hospital in Guadalajara showed no apparent broken bones, but judging by the 
amount of pain and the routine swelling by the end of the day, John felt that it 
was more than just a sprain. However, with my expertise in speaking Spanish, | 
managed to find ice at motels to treat the swelling. We were able to continue 
with our collecting trip. 

John was always the teacher. To pass the time when driving, we would grill 
each other on botanical glossary terms and other aspects of botany. I had never 
taken an economic botany course, but I survived the equivalent of one after 
experiencing the tropics with him. He was the opposite of me, since | eat to live 


SIDA 22(1): 51 — 54. 2006 


52 BRIT.ORG/SIDA 22(1) 


John Thieret, Chihuahuan Desert, Mexico, April 1970. 


— 


and he lived to eat. In the restaurants, meals were scrutinized very diligently 


ean 


and I was often quizzed on the content. He ate and sampled many kinds of fruits 
from the markets, many of which | had never heard of. In the field, he was al- 
ways tasting and smelling the fruits, leaves, etc. and discussing the results a 
spitting out the remains. His knowledge of tropical fruits was uncanny. 

We followed the coastline on our way home. As we neared the border, we 
drove past a bakery in a small town. John’s acute sense of smell overcame him 
and | had to turn around so he could stock up on some bread before we left 
Mexico. My common sense told me to refrain since we had come that far with 
no digestive problems. As it turned out, he did get a light case of diarrhea, but | 
did not pursue the issue. 


Ss 


ter 


At Mazatlan, we bought more newspapers (by the kilos) for our collections 
at the local newspaper office, and we spent our last night in Mexico south of 
Nogales, Arizona. U.S. Customs wanted time to go through our specimens look- 
ing for rusts for one of their projects. We went to dinner Cunch) and when we 
returned, customs had just finished. We headed back to Texas with our many 
collections and John’s foot still in pain. 

Once we got on the road, John got anxious to get home and check out his 
injury. We drove all night and stopped in Abilene, Texas for gas the next morn- 
ing. At Love Field in Dallas, I let him out at the airport where he caught a plane 


MAHLER, JOHN W. THIERET, COLLEAGUE AND ED 53 


John Thieret, Chihuahuan Desert, Mexico, April 1970. 


i 


and I went on home. I arrived home about five o'clock in the afternoon after 
being up and at the wheel for thirty-six hours. John told me later that they had 
found a hairline crack in one of his ankle bones in addition to the sprain. 
Although John was not one for attending meetings, he agreed to accom- 
pany me in the red Ford pickup to the Southwestern Association of Naturalists 
meeting in Tucson, Arizona. During registration, he headed over toa young man 
and shook hands. The fellow seemed confused and asked John if they had met 
before. John informed him that they had been officemates for ten years back in 
Chicago. John’s appearance was much different because he had shaved his beard 
that he had always had. Everyone had a good laugh about it, especially John. 
The field trips were productive and entertaining, but John’s most valuable 
contribution to my career was his involvement in Sida. Lloyd Shinners had told 
me that I needed to meet John when I first arrived at SMU, and he was right. 
When I inherited Sida, he, along with others, encouraged me to continue its 
publication, as opposed to others who felt | should drop the journal. With the 
help of Mrs. Helen Koresh, a sister of Lloyd's, | was able to clear the financial 
obligations Lloyd had encumbered with the authors of unpublished manu- 
scripts. [sent John some articles for him to review that I had written about Lloyd. 
John really “cleaned them up’ and returned them with more red ink than black. 


54 BRIT.ORG/SIDA 22(1) 


This was the beginning of his “internship” and status as Associate Editor of 
Sida that he held until his death. 

| have always considered myself lucky to have associated with many indi- 
viduals whose knowledg 
furthered my career goals. John was one of those individuals. The wide range 


e and insight enriched my professional knowledge and 


of things I learned from John—both professionally and asa friend—will always 
figure significantly in my lile. 


JOHN W. THIERET, 
ASSOCIATE EDITOR OF SIDA (1972-2005) 
Barney Lipscomb 
Botanical Researcl h Institute of Texas 


509 Pecan Street 
Fort Worth, Texas 76102-4060, U.S.A. 


John W. Thieret was an extraordinary and innovative teacher who shared his 
knowledge freely. Without doubt he was an influential botanist and one of the 
most distinguished editors of the 20th century at least in North America. “Big 
John,” as sometimes called him, was my steady and faithful cheerleader, Sida 
supporter, and crusader extraordinaire. He was always there whenever I needed 
advice or help with Sida or for that matter anything. Notwithstanding John’s 
many obligations, he called me after receiving each and every issue of Sida. 
And the voice I heard each time was inspirational, always filled with praise for 
the good but soft and tender on the good that could be better. John’s source of 
inspiration was perhaps from the many years of editorial experience with Eco- 
nomic Botany, Flora of North America, and the Journal of the Kentucky Acad- 
emy of Science. John knew first hand the amount of work involved and maybe 
sensed there was always room fora little more praise than was given [to editors 
everywhere]. “Keep up the fine work good ole fellow! How are you doing? How 
are you holding up? Let’s go on a field trip! Now is there anything you would 
like for me to do? Edit a manuscript? Review a book and write a review? What- 
ever it is just send it my way and [ll promptly return it to you.” John Thieret 
was forever supportive and never failed to do what he said he would do. He was 
always thinking of how he could help the other person and I always felt like | 
was that other person. John so loved Sida that he gave nearly 40 years of his life. 
To John be the glory—great things he has done for Sida, Contributions to Botany! 
This is my tribute to him. 


— 


Highlights of John Thieret’s contributions to Sida 

Ralph Thompson's (2006) excellent tribute provides a detailed account of the 
great service John Thieret rendered to botany and academia, his chosen fields. 
My tribute will recount his great and lasting contributions toa journal he cared 
so much about, Sida, Contributions to Botany. John’s love and early relationship 
with Sida was tied closely to that of his colleague and friend Lloyd H. Shinners 
at Southern Methodist University. Therefore, I pick up John’s trail of communi- 
cation with Shinners even before Shinners started Sida in 1962. 
1961—According to Ginsburg (2002), “Thieret was on staff of the Chicago Natural 
History Museum when he wrote to Shinners in 1961 to inquire about academic 


SIDA 22(1): 55 — 63. 2006 


56 BRIT.ORG/SIDA 22(1) 


positions for botanists. Shinners was very helpful, and when Thieret moved to 
the University of Southwestern Louisiana [Lafayette] the two became friends 
and close collaborators.” Thieret wrote, 

“I really think that a private line ought to be established between your herbarium and ours...” 


Music was another shared enthusiasm. 


1962—L loyd H. Shinners alone wrote, edited, and published the first two issues 
of Sida which were distributed on November 23rd. He said that he would have 
to publish his own journal because his own articles would never be published 
otherwise. So it was quite a tribute when he published others’ papers. 
1963—With the publication of the third issue of Sida on December 13, John 
Thieret became the fourth author aside from Shinners to have a paper published 
in the journal, behind W. Phillip Osborne, Walter H. Lewis, and Arthur 
Cronquist. John’s 54 page Yellowknife Highway paper followed Arthur 
Cronquist’s paper, The taxonomic significance of evolutionary parallelism. 


— 


Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. |: cata- 
logue of the flora. Sida 1(3):117-170. 


1964—Five of John’s manuscripts were submitted and published in 1964. The 
first issue contained the second part of John’s botanical survey along the 
Yellowknife Highway. In addition, Sida 1, no. 4 contained another significant 
and memorable event for John Thieret. A new flowering plant species was named 
in his honor for the first time ever. Lloyd Shinners (1964) named Scutellaria 
thieretii (Labiatae) for John. Lloyd wrote, “The Louisiana plant is considered to 
be one more in a group of very closely related species, and in honor of an ener- 
getic and productive collector it is named... am indebted to Dr. Thieret for sup- 
plying the two SMU collections, and for the loan of mounted specimens from 
the University of Southwestern Louisiana.” And so the relationship between 
John and Lloyd strengthens. 


Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. Il: veg- 
etation. Sida 1(4):187-239 


Shinners began work on Sida 1(5), but it was a struggle to get the manuscripts 
ready for the printer (Ginsburg 2002). Thieret apparently offered help with is- 
sue no. 5, but | could find no correspondence indicating whether or not Shinners 
ever took John up on his offer. Maybe John helped by submitting well-written 
and superbly edited manuscripts for Lloyd to publish. 


Fatoua villosa (Moraceae) in Louisiana: new to North America. Sida 1(4):248. 

Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) in Louisiana: new 
to the United States. Sida 1(4):249-250. 

More additions to the Louisiana flora. Sida 1(5):294-295. 


Eriogonum annuum (Polygonaceae): biennial in Nebraska. Sida 1(6):382. 


a 


LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 57 


1965—There was a break in the action for John as far as submitting papers to 
Sida. John did not have a single paper in either of the two Sida issues published 
in 1965. Regardless, John apparently stayed in close contact with Lloyd. 


pal 


1966—The next documented account of cor 
Lloyd took place on January 26. 


I ce between John and 
“Lam sending, for possible publication in SIDA, a manuscript entitled “Seeds of Some United States 
Phytolaccaceae and Aizoaceae.” The article was published on September 27. 


Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2:352—360. 


Sometime in 1966, Shinners had his first “all out” case of insulin shock and after 
hearing about that, Thieret wrote, “I was appalled to hear your account of the 
insulin shock.” I suspect John once again offered Lloyd help with Sida but ap- 
parently no letter with such words are in Shinners’ archives. Shinners recov- 
ered from the insulin attack and published a fourth issue of Sida in 1966: vol- 
ume 2, number 6 was published on December 30. 


1967—According to Ginsburg (2002) Shinners was probably exhausted after 
publishing four issues of Sida in 1966 and even more exhausted after his second 
NSF grant ended in May 1967. Thieret apparently kept in close enough contact 
to know the details. According to Ginsburg (2002), “He [Thieret] was willing to 
come to Dallas for a weekend to help in getting an issue together, or he would 
read and edit anything that Shinners m ight send.” Ina letter to Shinners, Thieret 
once again offered assistance with Sida. 


“Sida is a most worthy cause. To help out is a privilege...” 

John not only continued to correspond with Lloyd, but he also submitted manu- 

scripts to Sida and Lloyd published four of John’s papers in 1967. 

Supraspecific classification in the Scrophulariaceae: a review. Sida 3(2):87-106. 

Thirty additions to the Louisiana flora. Sida 3(3):123-127. 

Cyperus subgenus kyllinga (Cyperaceae) in the continental United States. Sida 3(3):128- 
136. [with AJ. Delahouhoussaye] 

Neogaerrhinum kelloggii (Greene) Thieret, comb. nov. (Scrophulariaceae). Sida 3(3):187, 


1968—John Thieret wrote Lloyd on April 18, 1968. 


Glad to hear, also, that SIDA is such a well-going concern. Once again I make an offer to come to 
Dallas for a weekend if I can be of help—real help—in getting together an issue. Also, I'll be glad to 
read and edit anything you send over. Even that Czech paper—which you specifically said not to edit, 


you will recall.” 
Two issues of Sida were published in 1968 but John did not have any papers in 
either issue. 


1969—Shinners published two issues of Sida in 1969: 3(5) and 3(6), August and 


58 BRIT.ORG/SIDA 22(1) 


November respectively. In between those two issues John sent Lloyd a letter 
dated September 9, 1969: 

“| was surprised to get the Bigelowia [Anderson] opus. Have you decided to establish an editorial board 
to “protect” your contributions against non-promotion? (Ea!) 

After hearing about another round of Shinners’ health problems, Thieret again 
offered to help proof read Sida. 

“You are willing | recall to have me take care of the index lor vol. 4 ol Sida. Right?” 

Meanwhile John submitted four manuscripts for publication. Undoubtedly they 
were submitted in excellent shape and Lloyd probably had to do very little work 
on these papers to get them ready for publication. ; 


Sagittaria guayanensis (Alismataceae) in Louisiana:new to the United States. Sida 3(6):445. 

Rumex obovatus and Rumex paraguayensis (Polygonaceae) in Louisiana: new to North 
America. Sida 3(6):445-446. 

actea (Rafinesque) Thieret, comb. nov. (Leguminosae). Sida 3(6):446. 


— 
— 


Baptisia 
Trifolium vesiculosum (Leguminosae) in Mississippi and Louisiana: new to North America. 
Sida 3(6):446-447. 

1969-1970—William FE Mahler was appointed three-quarters time assistant 
professor in the Biology Department at SMU and one-quarter time with the 
herbarium. Shinners’ Herbarium Report involved mostly Mahler's activities; 
Sida activities were still under the control of Shinners. Once again John offered 
Lloyd help in another letter probably dated sometime in April or May 1970 (see 
also Mahler's tribute). 

“Went ona collecting foray this past weekend—the first extensive one since I got back from Mexico. 
The ankle still gives trouble... Indeed, by the time the collecting trip was over, I was limping some- 
thing awful (). 

You were willing, I recall, to have me take care of the index for volume 4 of SIDA. Right???... If 
you are still, | shall get onto the task, a bit ata time. Shaw's paper alone is a challenge.” 
1971—Lloyd Shinners published Sida 4(2) on January 15, 1971 and one month 
later Shinners passed away on February 16, 1971. William F. Mahler inherited 
Sida from Lloyd’s sister Helen Koresh and he immediately became editor and 
publisher. February 17,1971, the very day after Lloyd’s death, John Thieret com- 
posed and mailed a letter to Mahler at SMU offering personal assistance and 
moral support. 


pele | 


“Just a note to reaffirm my offer to do what | can do to help keep SIDA a going thing. The journal 
a memorial to its founder. 


should certainly not be allowed to die but should be kept viable as, perhaps 
hel J; 
LF 


s for specific things could do, Icertainly would be glad ting. | would be willing 


to come over there every once in a while to work closely with you or with anyone else interested. 
Editing, reading proof, indexing—all would suit me fine. Speaking of indexing, Lloyd already gave 
me the go-ahead on doing the index for volume 4. 

Certainly you are thinking of an article on Lloyd, in SIDA, with possibly a list of his publica- 


tions. I'd be glad to help here, especially on the bibliographic work involved. 


LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 59 


eats yi offers b have ee made, It is now up to you to see that they are taken u 


ne to urge in to do all _ can to ee SIDA going?’ 


John Thieret was concerned about the future of Sida. Here is his letter to SMU 
urging Southern Methodist University’s President Dr. Willis M. Tate and the Dean 
of Graduate School, Dr. CC. Albritton, Jr, to continue the publication of Sida. 
“The name of the president I got from the 1969-70 edition of the WORLD OF LEARNING. Iassume 
that ie is ae the man. The Albritton part is for certain OK 


1 ] 


iat botan I sht (or something like that) and that I strongly 


urge SMU to aor all te can to continue the Seat of SIDA, a journal that is an excellent one that 
has filled a need. Etc. Etc. Ete. Signed John T. 


Even with all the encouragement from a number of Lloyd's colleagues, South- 
ern Methodist University never committed to preserving the journal or the col- 
lections that really made a name for the university in the botanical sciences. 
Other scientific journals were also born there but either died there or moved on 
to more supportive organizations. The dedication and hard work that Lloyd 
Shinners devoted in championing the cause of botany at SMU was kept alive 
through colleagues and the Botanical Research Institute of Texas (BRIT). Will- 
iam Mahler committed 21 years of personal time and money in keeping Sida, 
Contributions to Botany successful. In 1987, Mahler along with colleagues Lip- 
scomb and Andrea McFadden were instrumental in the establishment of the 
free-standing research institution BRIT. The core of the BRIT collections is the 
Lloyd H. Shinners Collection in Systematic Botany, originally at Southern Meth- 
odist University. Since Sida was privately published by Mahler it too joined BRIT. 
Mahler officially transferred ownership of Sida to BRIT in 1993. 

Mahler (1971) published Sida 4(3) with a tribute to Lloyd and John had 
one paper in the same issue: 


Physalis lagascae (Solanaceae) in Louisiana: new to the conterminous United States. Sida 
ADT F 


John’s offer of help was at last accepted. Mahler officially asked him to serve as 
associate editor of Sida in late 1971 or early 1972. One of the first papers John 
edited as associate editor was a large manuscript on the genus Rorippa sent to 
Dr. Shinners (early 1970) by Dr. Ronald L. Stuckey, associate professor of botany 
at Ohio State University. Dr. Stuckey expressed a desire to help with Sida ina 
letter dated February 24, 1971 to Dr. Mahler and Dr. Thieret. 

It has come to my attention that Professor Lloyd Shinners has passed away. I talked with John Thieret 
er]. My interest is that over a year ago I 


2 


by phone soy and he suggested that I write to you[Ma 
sent my d rtation, a sae aS peers on ties eeu ion apa to Dr. Shiners for publi- 


cation in Sida, which he ag nadditio enses for publi- 


cation, and this d to Sida i f $250.00 each, one in eee 1969 and one 
in December 1970. aie I would hope that you and Dr. Thieret, or anyone else, could see to it that the 
present manuscript on hand and/or in press for Sida could be published, and certainly as a memo- 


60 BRIT.ORG/SIDA 22(1) 


rial to Dr Shinners. | would be willing to do most anything, at this distance, to help tl se al 

| suspect financing is the big problem, and | would be willing to contribute some more funds, if | 
knew for sure my eee Male be published in Sida. Furthermore, I suspect some editing or 
deletions of my | dhe 


ge, 


elp. It was my understanding that the specimen citations would 


=~ 


not be pu plished. 
t also say that Mr. Robert Haynes, our assistant curator of the Herbarium, and former 
student of Dr. Thieret, has his manuscript on the genus Conopholis being published in Sida. 
[ would also like to see this fine journal for taxonomic botany and related interests be contin- 
ued either by Southern Methodist University or by a federation of the south-central botanists, for 
example. 


Mahler's letter (3 Mar 1971) back to Dr. Stuckey and again Dr. Thieret is involved: 


“In regard to your letter of 24 February 1971, we are planning to continue ea and follow Dr. Shinner’s 
plants for Vol. 4 (3) and (4). Issues 3 and 4 will be the paper lescribed in his letter of 26 March 
1970. 


You are being credited with a $500 contribution towards the publication of your MS. At this 
time, | cannot say what the financia 


— 


situation will be in relation to SIDA and its $12 page costs (re 
Shinners’ letter, 9 December 1969), but hopefully we can continue as leas by Dr. Shinners. 
My continuing to work like Dr. Shinners did would be impossible with the other activities (du- 


ties). He did not get a chance to go over the MS. in detail and I would like to send it toa reviewer such 
as Dr. Thieret. Perhaps you would like to make the changes Dr. Shinners suggested first, and then 
send it to Dr. Thieret. 

We have plenty of time in this respect as | do not expect No. 3 to come our till summer (71) at 
this time. 


[appreciate the briefing (previous correspondence) as it has been very helpful. | wish that ev- 
eryone would do it this way. Perhaps they will as I have one other individual who has brought me up 
to date in the same manner. 


The Conopholis paper of Haynes is still in the mill and status at this time unknown.” 


Dr. Stuckey’s response: 


Lenclose Xerox ¢ sof two letters which show that Dr. Reed C. Rollins has read and reviewed my 

Rorippa manuscript. | am eae that you have selected Dr. Thieret to go over my manuscript in 
' g 

preparation for its publication. It would be best if 


you transmitted the manuscript directly to him, 

rather than sending it back to me. The revision that Dr. Shinners mentioned involves a small item of 
about one page. | can do this and send the revision to Dr. Thieret. As you may know Dr. Thieret will be 
at our University for 5 weeks this summer teaching at Stone Laboratory. | will be there at the same 
time and can wor 


< with him on the manuscript at that time.” 
Mahler's letter (23 Mar 1971) to Dr. Stuckey and Dr. Thieret is mentioned: 


found your MS. yesterday (sti 


[have not located the plates, but will in time) and received your letter 
today. Dr. Shinners had discussed this MS. with me and stated that it would have to be trimmed to ca. 
the length of Kral’s paper 9ca. 175 pp.). As suggested by Shinners to you, ca. 150 pp. can be eliminatec 
readily (the citation of specimens). That still leaves ca. 100 pp. (70 pp. of MS. + 30 maps, illus.) to be 
eliminated. Thisi is one of the items of “reviewing” that | was referring to in regard to John Thieret. If 

u wish to do this yourself and possibly you should as it’s your MS., John Thieret might be helpful in 
short cuts so that you will not leave out any essential ena) The discussions, etc., are rather ver- 
bose and the same ideas could be expressed in fewer words.’ 


a 


1972—Thieret’s letter January 11, 1972) to Dr. Stuckey announcing his associ- 
ate editorship: 


LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 61 


SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. TAXONOMY AND DISTRIBUTION OF THE 
Mahler, SMU Herbarium, Dallas, Texas, 75222, U.S.A. Subscription price $8 GE es RORIPPA (CRUCIFERAE) 
(U.S.) per volume 


NORTH AMERICA 


RONAL DL. Such EY 


Columbus, Ohio 43210 


Rorippa consists of those yellow-petaled, numerous-seeded, readily- 
dehiscent-fruited Ley and shore “plans in the Cruciferae Distributed 
Berea in temperate the genus 

ccurs every continent except Antarctica, but the Species are more 


iber may increase to 60 or 70 
an expanded knowledge Tonal “(a968) jeanatderedtt here to be about 70 
species, some imperfectly known. One ea eeue Sees, R. palustris, 
is known fom eac me continent where t . Busch (1915) a 

Eurasia; s Warncinaencs (1960) ha 
eight species for Middle Europe; and, in Flora Europaea, yea “uss 
included 10 specie: 


ASSOCIATE EDITOR and So America. I 
7 cal ones) as native to North America; seven species are con: nsidered as 
John W. Thieret 
itoce The evolutionary relationship of these foreign members to the 


orth America, Rorippa is most common and abundant, both as to 
inva and t 


F 
aiene Arte irs of lakes te ce banks rea and 
rivers, and in marshes bee swales; or in wet inca Pinte sites 
ong roadside, drainage, and irrigation ditches 
in ni fallow or cultivated fields and gardens, along railroad grades and 
ditche: 


HISTORY OF THE GENUS 


rippa begins 
with Scopoli (Fi Carn, ed. 1, 520. 1760), who wrote the first, but brief, 
eneric diagnosi three species were 
named. The first contains the exact phraseology as that provided by Lin- 
naeus (Sp. Pl. 675. 1753) for attire sylvestre. Scopoli cited ee 
as one of five sources; on this basis, Rorippa sylvestris (Linnaeus) Besser 
s taken as the type of the genus. Abrams (1944) listed eae am- 


pat il fulfillment of the 
of Michipina Aaa 


SIDA Contributions to Botany, Volume 4 Number 4, pages 279—430 
Cop. >) aa The Herbarium 


ight 1972 
by wm, F. Mahler 


Left: Copyright page of Sida 4(4), the first issue of Sida in which John Thieret was appointed associated editor. Right: 
ope £D lac. I J. Davt co Look e.g 


rr 


“Thanks for l our Rorippa ms. Mahler is really the one to talk 
to about any financial matters. When I finish with the ms., | shall return it to Mahler. From then on, 
any dealings will be conducted by him. I have just been asked to serve as Associate Editor of Sida— 
which is just about what I've been doing all along.” 


John Thieret came to the editorial rescue as associate editor of Sida in 1972 and 
for the next 34 years, John proudly and faithfully served the journal in so many 
different ways. Mahler along with John’s superb editorial knowledge and skills 
and the help of many other colleagues, kept Sida going strong. Mahler passed 
the editorial baton to Lipscomb in 1983 but remained publisher until 1993; John 
was associate editor from 1972-2005. 


1972-2005—Not only was John an active associate editor of Sida, he was also 
contributing manuscripts. Twenty-seven of John’s papers were published in Sida 
during his 34 years of associate editorship. This represents almost 25% of all 
the papers John published in that same time frame. In fact, of all the scientific 
papers listed in Ralph Thompson’s tribute (2006), 26% were published in Sida, 
more than three times that of the next closest journal. The amount of work he 
contributed is enormous and the success, the foundation that John Thieret (and 
Wm. F. Mahler) made to Sida cannot be underestimated. We all have been for- 


62 BRIT.ORG/SIDA 22(1) 


tunate to have had John’s involvement with the journal but my (Lipscomb) ca- 
reer was truly impacted by this good friend. 
John’s influence on my (Lipscomb) personal career 
In the fall of 1975 my botanical career began as herbarium botanist at Southern 
Methodist University. There I worked in the great herbarium and botanical li- 
brary built by the prolific researcher, writer and founder of Sida, Lloyd Shinners. 
My first two years of work at SMU were strictly herbarium related and | casu- 
ally observed the editorial work on Sida by Wm. F Mahler. In the beginning I 
did not know who John Thieret was but it didn’t take me long to find out. Tra- 
dition is wonderful. Shinners told Mahler he needed to meet John and Mahler 
told me I needed to meet John. In 1977, two years into my job at SMU, I was 
appointed assistant editor of Sida by my mentor, friend, and colleague Bill 
Mahler. It was around this time—pre-email days—that | first began to commu- 
nicate with “Big John” Thieret over the phone and through letters. Over time | 
began to interact with him more and more on Sida’s issues and editorial mat- 
ters. It would be years later before | actually met the man behind the red pen. | 
had no idea John was the master of editing. | was too young to envision the 
immediate and future impact this one man would have on my life. He genu- 
inely cared for me as an individual and shared his expertise and knowledge 
with me freely. Through John, a whole new world of wisdom and knowledge 
was available at my doorstep. John Thieret took me in almost as if | was one of 
his own children. He (and Mahler) helped me to learn the role of an editor one 
red pen at a time. 

A few years passed before | finally got to meet John in person. He was ev- 
erything that I envisioned. | enjoyed one full week with John and his lovely 
wife Mildred back in the late 1990s. | stayed in their home in Alexandria not far 
from Northern Kentucky University (Highland Heights) where John worked. 
It was in their home that I had my first meal of tef (Eragrostis tef (Zucc.) Trot- 
ter). It wasalso at the Thieret home that John taught me the word “kex” or “kexes” 
for a dead stalk or stalks of herbaceous plants, “especially of large umbellifer- 
ous plants” (Oxford English Dictionary). | enjoyed four wonderful days with 
John traveling and botanizing through northern and eastern Kentucky. One of 
the many botanical things I learned from John on that trip was the edible fruits 
of Rhus typhina (Staghorn Sumac). He told me the drink from the sweet fruits 
was called “rhusade.” After returning to the Cincinnati area from out botanical 

field trip, John and I drove to one of his fun places to botanize called Jungle 
Jim’s International Farmers Market in Fairfield (Cincinnati), Ohio. One quote | 
found on the web said, “It is the ultimate food shopping adventure.” We walked 
into this humongous store and headed straight to the produce section and did 
we ever have a great time identifying plants and plant parts. It was an awesome 
experience with John, the economic botanist. And we visited another of John’s 


mo 


LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 63 


favorite plant places, Spring Grove Cemetery in Cincinnati. What an incredible 
place to study and learn about the many different woody plants and what a 
wonderful teacher I had. 

For 33 years John Thieret served Sida, Contributions to Botany as associate 
editor. His crusading editorial and scientific contributions to Sida are beyond 
measure! He was a good friend and advocate of Lloyd Shinners and Sida; he 
was a solid anchor of support to Wm. F Mahler in the storm of uncertainty that 
followed Lloyd's death; and he was a radiant lighthouse of editorial help to me 
as a young editor. He was the guiding light of editorial excellence and author- 
ity. Sida is today in large part because of John Thieret. His 33 years of editorship 
with Sida stands longer than anyone else associated with the journal. 

John Thieret was my rock and my anchor. His editorial energy and enthu- 
siasm and loyalty to Sida will forever be remembered and deeply appreciated! 
Did I ever say thank you John? I’m sure I did many times but let me say it in 
print in Sida one last time for everyone to see, Thank you John for all that you've 
done and continue to do through your teachings. 


Postscript: Changes in the journal, Sida no more 

I'm glad these tributes to John Thieret could be published in the journal that he 
loved so very much. Soon the journal Sida, Contributions to Botany will no longer 
be published by that name but instead will be called the Journal of Botanical 
Research Institute of Texas. The new journal will have a new look but generally 
the same format, botanical focus, and editorial direction will remain. The lead- 
ership of the Botanical Research Institute of Texas has decided the name “Sida” 
is offensive and has too much of a negative connotation because of its similar- 
ity to the word meaning AIDS in Spanish, French, Italian, and Portuguese. Per- 
haps the name change is a good thing but I am personally saddened to see 
Shinners’ original and long-standing name displaced. 


ACKNOWLEDGMENTS 


Ithank Gary Jennings (library /archive help), Ruth Ginsburg (archive help and 
reading the ms), and Ralph Thompson and Guy Nesom for helpful comments. 


REFERENCES 


BoTANICAL RESEARCH INSTITUTE OF TEXAS ArcHives. 2006. Lloyd Herbert Shinners archive. Botanical 
Research Institute of Texas library, Fort Worth. 

Ginspurc, R. 2002. Lloyd Herbert Shinners: by himself. Sida, Bot. Misc. 22 

Matter, W.F. 1971.Lloyd Herbert Shinners 1918-1971. Sida 4:228-231. 

SHINNERS, L.H. 1964. Scutellaria thieretii (Labiatae), a new species from coastal Louisiana. 
Sida 1(4):251-252. 

THompson, R.L. 2006. John W. Thieret (1926-2005). Sida 21:3-19. 


A NEW NARROWLY ENDEMIC SPECIES OF CLEMATIS 
(RANUNCULACEAE: SUBGENUS VIORNA) 
FROM NORTHEASTERN TEXAS 


Dwayne Estes 


abe of Tennessee Heal ium CERN 
Department of Ecology and Evolutionary Biology 
vines of Tennessee 


Knoxville Tennessee 37996, U.S.A. 


tnplants@yahoo.com 
ABSTRACT 
Clematis carrizoensis, a new ese! endemic species of subgenus Viorna, is described from the 
Carrizo Sands of northeastern Texas. This species belongs to a complex of similar species character- 
ized by glaucous and glabrous oe leaf surfaces and stems. Four additional species comprise this 
com ee C. addisonii, C glaucophylla C. texensis. and o versicolor denen carriZoensis 1s iS LOED NO: 
logi ically LLLUSL similar | to bluist it ] 


C. versicolor. It differs from those two coe in having ei cream ee lightly lilac- need se- 
pals. From C. addisoniiand C. glaucophylla, C. carrizoensis differs in sepal color and in havi = thicker 
and more reticulate peaves: All ns species of the complex are allopatric and occupy somewhat dif- 
ferent habitat Further discussion of their See eee 


differences, Poreene a@iabaren ane habitat preferences is provided. 


RESUMEN 


Se describe Cl ti Oe endo a de Carrizo Sands 


L oO 
pl ] 


en el noreste de Texas. Esta especie pert 
tener los tallos y el envés glaucos y glbros FI | complejo tae tras Caddicantt C 
glaucophylla, C. texensis, y C. versicolor. rizoensis es la mas fe ee a 
. Lees ep los escarlata ie C. ve mei sépalos que van del rojo purpura al lavanda azulado. 
ener los sépalos amarillo crema ligeramente tenidos de lila. De C. 
addisonii y C. aaeoonsi ite. se eee C. carrizoensis por el color de los sépalos y por tener las 
jas mas ee y mas Saaeconee . as cinco cece coBpSe son ope acaS y FOSpan 


morfoldgicas, Giseibucion peop y preferencias de habitat. 


In North America, 18 species of Clematis (Ranunculaceae) belong to the subge- 
nus Viorna (Pringle 1997). Of these, eight currently recognized species were 
included in the formerly recognized Clematis section Viorna subsection Viornae 
(Erickson 1943; Dennis 1976; Kral 1987): C. addisonii Britton, C. glaucophylla 
Small, C. morefieldii Kral, C. pitcheri Torr. & A. Gray, C. reticulata Walter, C. 
texensis Buckley, C. versicolor Small, and C. viorna L. These species are peren- 
nial vines characterized by relatively large (1-3 cm) urceolate flowers with thick 
leathery sepals that are mostly arranged in solitary or few-flowered cymes borne 


SIDA 22(1): 65 — 77. 2006 


66 BRIT.ORG/SIDA 22(1) 


on axillary peduncles subtended by leafy bracts (Dennis 1976). These taxa can 
be delineated into two groups based on leaf vestiture (Dennis 1976). The first 
group including C. morefieldii, C. pitcheri, C. reticulata, and C. viorna all have 
variously pubescent abaxial leaf surfaces and stems. The species of the second 
group, characterized by glabrous and glaucous abaxial leaf surfaces and stems, 
include C. addisonii, C. glaucophylla, C. texensis, and C. versicolor. These species 
form a complex here referred to as the C. glaucophylla complex. 

The species of the C. glaucophylla complex can be distinguished from each 
other by relatively few qualitative morphological characteristics. Often, a com- 
bination of features, including morphology, habitat, and distribution, must be 
used to identify taxa. Several structures including the caudex, roots, inflores- 
cences, and achenes offer little or no taxonomic value in separating most spe- 
cies of the complex (Erickson 1943; Dennis 1976). The most taxonomically use- 
ful traits are leaf texture, degree of leaf vein reticulation, and sepal color, 
although even these characters exhibit considerable variation and intergrada- 
tion (Erickson 1943: Dennis 1976). For instance, leaf thickness is useful when 
extremes are exhibited, but Dennis (1976) noted that factors such as leaf age 
and environmental conditions often make it a difficult character to assess. Sim1- 
larly, degree of vein reticulation is a useful character when found in its extremes 
but it can be confounding when trying to evaluate the continuum between ex- 
tremes (Dennis 1976). Erickson (1943) concurred, noting that while reticulate 
venation is a valuable taxonomic character, “it is not often possible to distin- 
guish between degrees of reticulation consistently.” Dennis (1976) considerec 
sepal color to be important for distinguishing species of the complex; for ex- 
ample thin-leaved specimens of C. versicolor can only be distinguished from C. 
glaucophylla by sepal color. Other characteristics, including degree of leaf dis- 
section, leaflet shape, flower shape, and stamen vestiture, are only generally 
useful for distinguishing species and are apparently never diagnostic. While 
the species of the C.glaucophylla complex may be difficult to differentiate mor- 
phologically, all members of the complex exhibit specific physiographic pref- 
erences (Dennis 1976) and have non-overlapping ranges (Fig. 1). 

Clematis texensisand C. versicolor are completely allopatric with the former 
restricted to the Edward's Plateau of Texas and the latter confined to the Inte- 
rior Highlands (Arkansas, Missouri, Oklahoma) and Interior Low Plateaus (Ala- 
yama, Kentucky, Tennessee). They differ consistently in flower color and gen- 
erally in leaf dissection and leaflet shape. Both species usually have relatively 
thick and conspicuously reticulate leaves. Erickson (1943) called attention to 
possible intermediate populations occurring in the region between the Interior 
Highlands and the Edward's Plateau noting that some specimens from this re- 
gion have been difficult to assign to a particular species. Erickson arbitrarily 
referred questionable specimens from southwestern Arkansas and southeast- 
ern Oklahoma to C. versicolor and specimens from northeastern Texas (Smith 


— 


en 


ees 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 67 


Legend 
Clematis spp. 

@ = §=C. addisonii 

@ = $C. carrizoensis 
= ~=6C. glaucophylla 
C. texensis 


4 ~~ C. versicolor 


Co TkKilometers 


Fic. 1.G hi f the fi i ftha Cf tie al hull | fr aera ) i 1976) 


County) to C. texensis. Dennis (1976) contended that the Texas specimens were 
not C. texensis because the sepals were white to pale lavender rather than the 
bright red characteristic of that species. Interestingly, Dennis (1976) referred 
the Texas plants to C. versicolor in spite of the fact that they also differed from 
that species in sepal color and habitat and occurred ina different physiographic 
region ca. 200 km from the nearest known C. versicolor population. Additiona 

study of herbarium specimens from this region coupled with field work has 
revealed that the Texas populations previously referred to C. versicoloractually 
represent a new species described below. 


Clematis carrizoensis D. Estes, sp. nov. (Figs. 2, 3, 4B). Type: U.S.A. TEXAS: Smith Co. 1.5 
mi NW of Lindale, sandy soil, vine growing along fence near roadside, 15 May 1949, ].F Hennen 
408, consists of 2 sheets (HOLOTYPE: SMU). 


jen 


A Clematide versicolore sepalis e lilacinis flavidis-cremeis, foliis plerumque dissectioribus, foliolis 
saepiuscule emarginatis ad apices profundius cordatis basibus differt. 


Perennial trailing or scandent vines to 3 m long, from a woody caudex bearing 
numerous smooth fleshy apparently unbranched roots. Stems stout proximally, 
slightly flexuous distally, 6 angled, 12 ribbed, glabrous and glaucous, reddish- 
brown to pale brown. Leaves oppositely decussate, imparipinnate to pinnate- 


68 BRIT.ORG/SIDA 22(1) 


THE UNIVERSITY OF TENNESSEE 


1 Small 


W. Michael Dennis December , 476 


PLANTS OF TEXAS 


Clematis reo ulata Valte 


| HOLOTYPE OF: 
| 


Smith Co.1 1.5 miles nortivest of Lindale. 


Sanly soil. Vine growing along; fence 
near roadside. 


| Ctesniit ; sis D. Estes 


Sheet 1 of 2 J. Ps Hennen 1,08 May 159 1oL9 


Fic. 2. Holotype (UJ. Hennen 408, SMU) of Clematis carrizoensis. 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 69 


Fic.3. Clemati ji is. A. Portion of plant showing g| baxial leaf surfaces. B. Close-up of abaxial surface of 
| ins). D. Fruit. Photos above 


leaflet showing raised-reticulate venation. C. Flower, fi iew ( isy I 
f | H] 1 i D. Estes 07032 


P 


ternate, largest at about mid-stem, spreading, to 2.0 dm long, longer than wide 
to as wide as long; petiole stout, equaling or more frequently shorter than the 
lowest leaflets, to 6.0 cm long, glabrous and glaucous, base of opposing petioles 
usually connate; rachis geniculate; leaflets symmetrical or asymmetrical, 
paired, 8-10 on imparipinnate leaves, to 25 on proximal-most pinnate-ternate 
leaves, spreading to erect, reduced distally on rachis, the distal-most often form- 


70 BRIT.ORG/SIDA 22(1) 


ing a short simple tendril, proximal-most entire or 2-3-lobed to trifoliate, ovate, 
broadly ovate, or subrotund, leaflets to 7.0 cm long and to 6.0 cm wide, coria- 
ceous, base shallowly to deeply cordate, rounded, or truncate, margins slightly 
revolute, leaflet apices and lobe apices rounded, retuse, or occasionally acute, 
mucronate, adaxial and abaxial surlaces strongly reticulate with all vein or- 
ders conspicuously raised, forming an intricate reticulum, light green adaxially, 
paler abaxially, drying pale olive-brown to olive-green, glabrous and glaucous 
on both surfaces but more conspicuous on abaxial surface; petiolules 1.5-3.0(-7 
cm) long, one-fourth to one-third length of leaflet or rarely one-half to as long 
as leaflet, stout, glabrous and glaucous. Flowers solitary on long axillary pe- 
duncles or rarely axillary and terminal, nodding, ovoid or urceolate; peduncles 
erect, curved or frequently abruptly recurved at apex, 7.5-17.0 cm long, stout, 
glabrous and glaucous, with 2 foliose bracts; bracts sessile or subsessile with 
petioles to 4.0 mm, positioned mostly near the middle of the peduncle, base 
subcordate to deeply cordate, margins entire or 2-3-lobed, otherwise similar in 

shape, size, texture, and vestiture to leaflets. Sepals erect, connivent, ovate-lan- 
ceolate to oblong-ovate, apices reflexed, 1.8-3.0 cm long, 0.7-1.3 cm wide, coria- 
ceous, ca. 12 nerved, abaxially yellowish-cream, or greenish-yellow and often 
faintly tinged with lilac proximally, adaxial surface similar in color to the 
abaxial surface at least distally, mostly drying tan or light brown, edges nar- 
rowly crispate, white-tomentulose, abaxial and adaxial surfaces glabrous. Sta- 
mens linear, 1.8-2.4 cm long, the filaments flattened, in dried material dark 
brownish-red in the proximal half, densely pilose from below middle to apex, 
anthers 6.0 mm long, connective extended 1.75-2.0 mm beyond anthers, pilose. 
Fruit tightly clustered in spherical heads 4.5-8 cm in diameter, light brown, 
body rhomboidal-ovate, 6.0-9.0 mm long and 6.0-7.0 mm wide, acuminate at 
both ends, compressed, marginally thickened, sericeous; style curvate, 3.0-5.5 
cm long, with a yellowish-brown plumose coma, hairs of coma spreading to 

scending, to 3.0 mm long. Chromosome number unknown. 


aoa specimens ene TEXAS: Cherokee Co.: ca. 9.5 mi E of Troup, ca.6 mi W ol Price, ca. 

ni N of Hwy 13, sandylands, 12 Jul 1977, G. Bee 50. 38 (BRIT). Smith Co.: sandy woods east ol 
wee 16 May 1902, J. Reverchon ee [two sheets] (MO); on R.R. [railroad] near Lindale, Apr 1923, J. 
Reverchon s.n. (MO); western Tyler along Cotton Belt Railway, east end of Pine Burr Rd., infrequent at 
edge of woods along railway, 21 May 1949, V.L. Cory 56212 (SMU); western Tyler, S of Pine Burr Rd, 
infrequent in sandy woods, 10 Aug 1950, V.L. Cory 57860 (SMU); just N of Interstate 20 in woods E ol 
Old Tyler-Mineola Rd., 20 Oct 1965, D.S. Correll & H.B. Correll 32041 (LL); sandy field with scattered 
post oak in SW angle where US Rte. 69 and Loop 323 intersect, north edge of Tyler, 28 Aug 1966, D.S. 
Correll 33513 (LL); pure sand in field on Loop 323 near intersection with Rte. 69, Tyler, 6 Jun 19609, D.S. 
Correll 37401 (LL); in SW angle of intersection of US 69 and Loop 323, N edge of Tyler, under oaks 


next to 12 Oaks Motor Hotel, growing in sandy field under Quercus incana, Q. margaretta, with Opun- 
tia, Yucca, 22 Jun 1975, WM. Dennis 5I718 (TENN); NW portion of city of Tyler, 0.75 km WSW of 
junction ol TX State Hwy 323 Loop and US Hwy 69, ca. 20 m SW of junction of Hwy 323 Loop and 

Silver Creek Dr, 32°22'42.1"N, 95°2020.6"W, along N edge of scrubby oak-hickory woods and in a 


small opening nearby, rooted in deep sand and scram acorer small shrubs and herbs, with Quercus 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 71 


\“}. 


spp., Carya, Rhus aromatica, Prunus gracilis, and Toxicodendron pubescens, 9 May 2005, D. Estes 07032 
(GH, NY, TENN, VDB). Van Zandt Co.: 2.3 mi E of Ben Wheeler, sandy loam, 6 May 1951, R. Van Vleet 
1257 (SMU). 


Distribution and Habitat.—Clematis carrizoensis is endemic or nearly endemic 
to the Carrizo Formation, a narrow zone (19 km wide) of relatively deep depos- 
its of coarse to find sand that trends southwest to northeast for ca.720 km from 
south-central to northeastern Texas (Macbryde 1933; Sorrie and Weakley 2001). 
Sorrie and Weakly (2001) identified an additional 12 taxa endemic or nearly 
endemic to this formation. Within the Carrizo Sands, C. carrizoensis has only 
been collected from three contiguous counties, Cherokee, Smith, and Van Zandt, 
all in the northeastern section of the formation. Clematis carrizoensis grows in 
or along the edges of open, well-drained, prairie-like areas in full sun or partial 
shade. Within these communities, C. carrizoensis trails along the ground or Over 
low herbs, shrubs, or fences, apparently rarely climbing higher into low 
branches of trees. Oak-hickory woodlands or thickets border these sites. Domi- 
nant tree species associated with one population of C. carrizoensis in Tyler, Texas 
(site of D. Estes07032 cited above) include Quercus incana Bartr., Q. marilandica 
Muench., 9. nigra L., Q. stellata Wang., Carya texana BuckL., Sassafras albidum 
(Nutt.) Nees, and Ulmus crassifolia Nutt. Frequent small trees, shrubs and woody 
vines were Opuntia sp., Prunus caroliniana (P. Mill.) Ait. P gracilis Engelm. & 
A. Gray, Rhus aromatica Ait. R.copallinum L., R.glabraL., Rubus trivialis Michx., 
Stillingia sylvatica Garden ex L., Toxicodendron pubescens P. Mill., Vaccinium 
arboreum Marsh., Viburnum rufidulum Raf., Vitis mustangensis Buckl., V. 
rotundifolia Michx.and Yucca sp. Baptisia sp., Cnidoscolus texanus(MuellL.-Arg.) 
Small, Croton argyranthemus Michx., Mimosa sp., Paronychia drummondii Torr. 
& A. Gray, Pteridium aquilinum (L.) Kuhn, Rumex hastatulus Baldw.,, 


— 


72 BRIT.ORG/SIDA 22(1) 


Schizachyrium scoparium (Michx.) Nash and Talinum rugospermum Holzinger 
were closely associated herbaceous taxa. 

Phenology.—Clematis carrizoensis flowers from early May to early August: 
fruits are produced beginning in late June and persist until November. 

Etymology.—The specific epithet “carrizoensis” was selected in honor of the 
Carrizo Sands, the geologic formation to which C. carrizoensis is apparently 
restricted. 

Common Names.—Carrizo sands leather-flower; sand clematis 

Conservation Significance.—Clematis carrizoensis is a rare species. | have 
seen only 12 specimens from three counties in northeastern Texas; 10 of these 
are from just four sites in a small area of Smith County. Label data from her- 
barium specimens also indicate that this species is uncommon where it occurs. 
Most of the specimens examined for this study were collected over 30 years ago 
possibly indicating that this species is declining or simply overlooked. Most of 
the populations represented by collections are located near or in the city of Tyler, 
Texas. These populations may be subject to extirpation by future development. 
This species should be searched for in other sandy sites, particularly those as- 
sociated with the Carrizo Formation, in northeastern Texas as well as in sandy 
habitats in nearby portions of Miller County, Arkansas and Caddo Parish, Loui- 
endangered and should be afforded 


KL 


siana. This species should be considerec 
protection at both the state and federal level. 


Species of the Clematis glaucophylla Complex 

Five species comprise the C. glaucophylla complex. While all species differ in 
rather subtle morphological characteristics, they are all allopatric and mostly 
inhabit different plant communities in different physiographic provinces. The 
complex is in need of a detailed phylogenetic study. 

Clematis addisonii, a narrow endemic of a four county area in the Ridge 
and Valley of western Virginia (Fig. LD), inhabits dry calcareous woods and out- 
crops (Kral 1983). Itis characterized by a mostly erect, non-twining habit, mostly 
simple, thin, and inconspicuously reticulate cauline leaves, and solitary termi- 
nal {lowers that have purple to reddish-purple sepals with whitish apices. 

Clematis glaucophylla ranges {rom northwestern Florida west to southeast- 
ern Mississippi and north to southeastern Tennessee (Fig. L); it is reportedly dis- 
junct to McCurtain County, Oklahoma (Dennis 1976). Reports of C.glaucophylla 
from Kentucky Jones 2005) are based on misidentified specimens (D. Estes, 
pers. obs.). Throughout most of its range in the Piedmont and Gulf Coastal Plain, 
C. glaucophylla grows in sandy neutral to slightly acidic soils along streams 
(Dennis 1976), but at some sites in Alabama and southeastern Tennessee, C. 
glaucophylla grows at the edges of dry rocky calcareous forests (D. Estes, pers. 
obs.). Clematis glaucophylla hasa oe habit and compound cauline leaves that 
have mostly thin and inconspicuously reticulate leaflets. The proximal leat- 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 73 


lets are often deeply lobed, divided, or ternate. Flowers of C. glaucophylla are 
borne on axillary peduncles and have sepals that are cherry-red or reddish- 
purple abaxially and light colored near the abaxial tip and on the whole sur- 
face adaxially. 

Clematis texensis is endemic to the Edward’s Plateau of Texas (Fig. 1) where 
itis often associated with calcareous woodlands, edges, cliffs, and stream banks. 
The species hasa viny habit and compound cauline leaves with relatively thick 
and conspicuously reticulate leaflets. The proximal leaflets vary from simple 
to lobed or ternate. The flowers are borne on axillary peduncles and the sepals 
are scarlet or rose-red abaxially and near the tip adaxially (Fig. 4). 

Clematis versicolor has a bicentric distribution with part of its range cen- 
tered in the Interior Highlands of southwestern Missouri, western Arkansas, 
and eastern Oklahoma and the other portion centered in the Interior Low Pla- 
teau of central Tennessee, extreme northern Alabama, and south-central Ken- 
tucky (Fig. 1). It is usually associated with calcareous bluffs, dry woods, and 
roadsides with exposed limestone. Clematis versicolor has a viny habit, relatively 
thick and conspicuously reticulate compound cauline leaves with the proxi- 
mal leaflets entire or rarely lobed or divided, and flowers borne on axillary 
peduncles. The abaxial surfaces of the sepals of C. versicolor are mostly pur- 
plish-red proximally and often are cream-colored or greenish distally (Fig. 4). 

Clematis carrizoensis is restricted to sandy prairie-like areas among oak- 
hickory woodlands in three counties in northeastern Texas (Fig. 1). The species 
hasa viny habit, thick and conspicuously reticulate compound leaves that have 
proximal leaflets frequently lobed, divided, or ternate, flowers borne on axil- 
lary peduncles, and sepals that are al lly yellowish-cream and faintly tinged 
with lilac proximally (Fig. 4). 


Characters Useful for Distinguishing Members of the Clematis glaucophylla 
Complex 

The members of the C. glaucophylla complex can be distinguished by differ- 
ences in habit, leaf dissection, leaf texture, leaflet shape, degree of vein reticu- 
lation, flower position, sepal color, habitat, and geography (Table 1; Fig. 1). 

In terms of habit, C. addisonii is the only erect non-viny member of the 
complex. It has stems that are mostly less than | m tall whereas the other spe- 
cies in the complex have viny stems mostly greater than 1 m in length. Dennis 
(1976) noted that while C. addisonii is the only species in the complex that ex- 
hibits this growth form, occasional individuals of C. addisonii bear apical 
branches that often become entangled and approach a viny habit. 

Leaf dissection is a character that is only generally useful for distinguish- 
ing species of the complex, the exception being C. addisonii. Clematis addisonii 
has leaves that are mostly simple but occasionally vary to pinnate with up to 
six leaflets. The rest of the species of the complex have leaves that are pinnate 


74 BRIT.ORG/SIDA 22(1) 


Taste 1.Characteristics of the Clematis glaucophylla complex, compiled from data pr ted in Dennis 
(1976) and Pringle (1997). 


texensis carizoensis versicolor addisonii glaucophylla 
iny>1m viny,> 1m viny,> 1m erect,< 1m viny,> 1m 
Leaf Dissection |-pinnate to ]-pinnate to 1-pinnate, simple, or distal 1-pinnate to 
pinnate pinnate- proximal leaves 1-pin- —_ pinnate- 
ternate, ternate, leaflets nate ternate, 
proximal leaf- — proximal leaf- ee proximal 
lets commonly lets commonly unlobec leaflets 
lobed lobed dk commonly 
or 3-foliate or 3-foliate 2-3 lobed lobed 
or 3-foliate 
Leaf Texture coriaceous coriaceous coriaceous thin thin 
Venation reticulate, reticulate, reticulate, not reticulate, — not reticulate, 
raised raised raised not raised not raised 


Flower Position mostly axillary mostly axillary mostly axillary mostly terminal mostly 


axillary 

Flower Color scarlet to yellowish pure redto purple to cherry red to 
e-red cream, ee os nder, purplish-red, — reddish- 
lilac-tinged creamy OF whitish distally purple 


proximally greenish distally 


to pinnate-ternate with more than six leaflets. The degree of compounding of 
the proximal leaflets is a useful yet non-diagnostic character that according to 
Erickson (1943) “must be used with judgment.” For example, Erickson (1943) 
noted that leaf dissection may generally be used to separate C. texensis and C. 
versicolor. Clematis glaucophylla, C. texensis,and C. carrizoensis frequently have 
lobed, divided, or ternate proximal leaflets whereas the proximal leaflets of C. 
versicolor are usually unlobed or rarely lobed or ternate. 

Erickson (1943) considered leaflet shape to be of general use in delimiting 
taxa, especially the leaflet apices. Generally, C. addisonii, C. glaucophylla, and 
C. versicolor have broadly to narrowly obtuse leaflet apices. Clematis texensis 
and C. carrizoensis have leaflet apices that are more frequently emarginatec 
apically but that vary to obtuse. Leaflets of C. carrizoensis, C. texensis, and C. 
versicolor are almost always mucronate at the apex. 

Among the species of the C. glaucophylla complex, two groups can be de- 
lineated based on leaf texture and degree of leaf vein reticulation. Clematis 
glaucophyllaand C. addisonii belong toa group characterized by relatively thin 
and non-reticulate leaves. The other group, consisting of C. carrizoensis, C. 


Wu 


~ 


texensis and C. versicolor, has relatively thick and conspicuously reticulate 
leaves. Once again, these characters are relative and are hard to differentiate 
when only one species is encountered and can sometimes be difficult to assess 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 75 


from herbarium material. Furthermore, shade plants of all species of the com- 
plex may be thin and non-reticulate. 

Flower position is a character that is of limited taxonomic value in the C. 
glaucophylla complex. Most species have flowers borne on axillary peduncles, 
however, C. addisonii has flowers that are mostly borne on terminal peduncles. 

Perhaps one of the most useful characters for distinguishing the members 
of the C. glaucophylla complex is sepal color (Table 1, Fig. +). Unfortunately, it is 
also the character that is most difficult to preserve and interpret on herbarium 
specimens. In C. glaucophylla, the sepals are more frequently cherry red to red- 
dish-purple abaxially except distally they are often light colored. Clematis 
texensis has sepals that are generally scarlet or rose-red, and the red coloration 
is present on both the abaxial surface and on the distal portion of the adaxial 
surface. Compared to C. glaucophylla and C. texensis, the sepals of C. addisonii 
are more purplish, varying from light purple to purplish-red with whitish api- 
ces. Typical specimens of C. versicolor have sepals that are purplish-red or blu- 
ish-lavender proximally and cream-colored or greenish distally. The sepals of 
C.carrizoensis are yellowish-cream and lightly tinged with lilac proximally. 

Habitat can be used to distinguish the species of the C. glaucophylla com- 
plex to some extent. Clematis addisonii, C. texensis, and C. versicolor are appar- 
ently obligate calciphiles. Clematis glaucophylla is a facultative calciphile. 
Clematis carrizoensis is not associated with calcareous rock outcrops but rather 
is restricted to deep sandy soils. All of the species seem to grow best in edge 
habitats: however C. addisonii, C. texensis, and C. versicolor are sometimes found 
in closed-canopy forests. 

When incomplete or non-flowering specimens of species of the C. 
glaucophylla complex are encountered, the most reliable character that can be 
used to distinguish the species is geography. All of the taxa are allopatric. The 
ranges of C. versicolorand C.glaucophylla abut one another in portions of south- 
eastern middle Tennessee and possibly extreme northern Alabama. Clematis 
texensisand C. carrizoensisare restricted to Texas and C. addisoniti is restricted 
to Virginia. Clematis versicolor is found only in portions of Oklahoma, Arkan- 
sas, Missouri, Kentucky, Tennessee, and Alabama; it should be also searched for 
in extreme northeastern Mississippi (Tishomingo Co.) and in extreme south- 
ern Illinois. Clematis glaucophylla occurs in Mississippi, Alabama, Georgia, 
Florida, and Tennessee. Alabama and Tennessee are the only two states that 
support more than one species of the complex. 


KEY TO THE SPECIES OF THE CLEMATIS GLAUCOPHYLLA COMPLEX 
(ADAPTED FROM ERICKSON 1943, DENNIS 1976, AND PRINGLE 1997) 
1. Plants ascending to erect, sometimes branched above, rarely viny; middle and lower 
cauline leaves mostly simple; flowers mostly terminal; restricted to Ridge and Valle 
Province of western Virginia C. addisonii Britton 


76 BRIT.ORG/SIDA 22(1) 


1. Plants scrambling or climbing, viny; middle and lower cauline leaves compound; 
lowers axillary; species not erviginles. 
2. Leaves usually thin,ad without raised secondary and ae veins 
C. glaucophylla Small 
2. Leaves usually thick, adaxial surface with moderately to prominently raised 
secondal y al id tertiary veins 
3. Abaxial surface of sepals scarlet to rose-red or PSOE C AID and one 
lilac tinged; leaflets mostly ovate to subrotund, base mostly subcordate to 
deeply cordate, apex usually Ue lower leaflets commonly 2-3- 
lobed or 3-foliate; plants restricted to Tex 
4. Sepals scarlet to rose-red abaxially; stamens not densely or uniformly 
pubescent, connective mostly glabrous or with a few stiff trichomes; 
plants of limestone uplands of the Edwards's Plateau in south-central 


Texas C. texensis Buckley 

4. Sepals yellowish-cream abaxially and lightly tinged with lilac; stamens 

densely and uniformly pubescent, connective moderately to densely 

pubescent; plants of deep sandy habitats of northeastern Texas __ C. carrizoensis 
Estes 


3. Abaxial surface of sepals purplish-red or bluish-lavender proximally and 
cream-colored or greenish distally; leaflets mostly ovate to elliptic, base 
obtuse to subcordate, apex mostly obtuse; lower leaflets mostly unlobed 
and undivided, infrequently lobed or ternate; plants of the Interior High- 
lands of Missouri, Oklahoma, and Arkansas and the Interior Low Plateau of 
Tennessee, Kentucky, and Alabama 


C. versicolor Smal! 
ACKNOWLEDGMENTS 


I would like to thank the faculty and staff at the following herbaria for loaning 
specimens or facilitating visits during this study: BRIT, MO, SMU, TENN, TEX- 
LL. Gratitude is also extended to R.L. Small, B.E. Wofford, and J. Beck for review- 
ing earlier versions of this article, to R. Naczi for preparing the Latin diagnosis, 
to J. Singhurst for assistance with field work, to C. Fleming for preparing the 
map, to S. Baskauf for use of his photo of C. versicolor,and to WC. Cart for use of 
his photo of C. texensis. 


REFERENCES 


Dennis, W.M.1976.A biosystematic study of Clematis, sect. Viorna, subsection Viornae PhD. 
Dissertation. The University of Tennessee. Knoxville 

Erickson, R.O. 1943. Taxonomy of Clematis section Viorna. Ann. Missouri Bot. Gard. 30:1-62. 

Jones, R.L. 2005. Plant life of Kentucky: an illustrated guide to the vascular flora. The Uni- 
versity Press of Kentucky, Lexington. 

Kral, R. 1983. A report on some rare, threatened, or endangered forest-related vascular 
plants of the South. USDA Forest Service, Southern Region, Tech. Publ, R8-TP2. 2 vol- 
umes. 


KRAL, R. 1987. A new “Viorna” Clematis from northern Alabama. Ann. Missouri Bot. Gard. 
74:665-669. 


ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 77 


McBryoe, J.B. 1933. The vegetation and habitat factors of the Carrizo sands. Ecol. Monogr 
3:249-297 

Prinate, J.S. 1997. Clematis. In: Flora of North America north of Mexico. Volume 3. 
Magnoliophyta: Magnoliidae and Hamamelidae, ed. Flora of North America Editorial 
Committee. Oxford University Press, NY. Pp. 158-176. 

Sorrie, B.A.and A.S.Weaktey. 2001. Coastal Plain vascular plant endemics: phytogeographic 
patterns. Castanea 66:50-82. 


78 BRIT.ORG/SIDA 22(1) 


ANNOUNCEMENT 
The Rupert Barneby Award 


The New York Botanical Garden is pleased to announce that Rodrigo Duno de 
Stefano, of the Centro de Investigacion Cientifica de Yucatan A.C. (CICY), is the 
recipient of the Rupert Barneby Award for the year 2006. He will be studying 
the family Leguminosae in the Yucatan Peninsula Biotic Province (YPBP), 
Mexico. With about 60 genera and more than 260 species there, the Leguminosae 
are one of the most important plant elements of the Yucatan region. This study 
will also contribute toa revision of four legume genera for the “Illustrated Flora 
of the Yucatan Peninsula” (G. Carnevali, general editor). 

The New York Botanical Garden now invites applications for the Rupert 
Barneby Award for the year 2007. The award of US$ 1,000.00 is to assist research- 
ers to visit The New York Botanical Garden to study the rich collection of 
Leguminosae. Anyone interested in applying for the award should submit their 
curriculum vitae, a detailed letter describing the project for which the award is 
sought, and the names of 2-3 referees. Travel to the NYBG should be planned 
for sometime in the year 2007. The application should be addressed to Dr. James 
L. Luteyn, Institute of Systematic Botany, The New York Botanical Garden, 200% 
Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA, and received no later 
than December |, 2006. Announcement of the recipient will be made by De- 
cember 15¢, 


Anyone interested in making a contribution to 
THE RUPERT BARNEBY FUND IN LEGUME SYSTEMATICS, 
which supports this award, may send their check, payable to 
e New York Botanical Garden 
Dr. James L. Luteyn 
Institute of Systematic Botany 
The New York Botanical Garden 
200th St. and Kazimiroff Blvd. 
Bronx, NY 10458-5126 USA 


SIDA 22(1): 78. 2006 


HILL CANE (ARUNDINARIA APPALACHIANA), 
A NEW SPECIES OF BAMBOO (POACEAE: BAMBUSOIDEAE) 
FROM THE SOUTHERN APPALACHIAN MOUNTAINS 


JK. Triplett! A.S.Weakley 


Department of Ecology, Evolution Univ. of North Carolina Herbarium (NCU) 
and Organismal Biology North Carolina Botanical Garden 
lowa State University University of North Carolina 


Ames, lowa, 50011-1020, U.S.A. Campus Box 3280 
Chapel Hill, North Carolina, 27599-3280, U.S.A. 
L.G. Clark 
Department of Ecology, Evolution 
and Organismal Biology 


lowa State University 
Ames, lowa, 50011-1020, U.S.A. 


ABSTRACT 


A newly recognized Sees of A Arundinaria from the southern Appalachian Moun ae! is nla 
illustrated, and compared with the related A.giganteaand A. tecta.A 

is distinguished by a combination of eecetiye morp hological characters hn eatures of 
branching ane as TORO By) a anatomy, and ecology. Recognition of this species is consistent 


monophy ly of the species and its sister relationship with 
A. te a A key for the eoaaienen of Arundinaria species in North America is included along with 
a comparative table based on morphology, leaf anatomy, and ecology. 


Key worps: bamboo, Arundinaria, eastern U.S., Appalachian Mountains 


RESUMEN 
— a ee pecie de Arundinaria del sur de la Cogulers ae ies saan y se 
; | : laci las A gigantea y A.tecta.S 
r z= S 
por una 1 aad g vegetati 70. I] P le | if 
morfologia y anatomia de las hojas, y ecologia F] a : A dar 


£ 
genéticos que apoyan la monofilia de la especie nueva y su posicion como especie hermanaa A. tecta. 
Se incluye una clave para la identificacion de especies de Arundinaria en América del Norte y una 
tabla comparativa basada en la morfologia, anatomia de las hojas, y ecologia. 


Pacapras claves: bambu, Arundinaria, este de los Estados Unidos, la Cordillera Appalachiana 


INTRODUCTION 
Arundinaria Michx. is a genus of north temperate woody bamboos (Poaceae: 
Bambusoideae) with a complex taxonomic history involving numerous enti- 
ties that have been placed within it at one time or another over the past two 


‘Author for correspondence (triplett@iastate.edu) 


SIDA 22(1): 79 - 95, 2006 


80 BRIT.ORG/SIDA 22(1) 


hundred years (McClure 1973; Li 1997; Judziewicz et al. 1999), with older treat- 
ments including upwards of 400 heterogeneous species from Asia, Africa, and 
the Americas. Currently Arundinaria is treated in a restricted sense to include 
only those species endemic to the eastern United States (Ohrnberger 1999), but 
debate continues regarding the inclusion of certain Asiatic taxa (e.g., Pleioblastus 
Nakai, Pseudosasa Makino ex Nakai, Bashania P.C. Keng & Yi, and 
Oligostachyum Z.P. Wang & G.H. Ye) that share key morphological features with 
the North American species (Li 1997; Judziewicz et al. 1999). However, because 
Arundinaria gigantea (Walter) Muhl. is the type species for the genus, its ge- 
neric placement is secure. As such, Arundinaria represents the only bamboos 
native to North America and the only temperate bamboos (the North Temper- 
ate clade of Clark et al., in press; Zhang & Clark 2000; Kelchner & Clark 1997) 
native to the New World, and provides another example of the classic disjunc- 
tion pattern in the flora of eastern Asia and eastern North America (Wen 1999). 
With a species richness ratio of approximately 20:1 up to 90:1, depending on 
which Asian group is used for comparison within the North Temperate bam- 
boo clade, Arundinaria and allies also provide an example of the intriguing 
asymmetry within this disjunction discussed by Guo and Ricklefs (2000). 
Arundinaria s.s. encompasses arborescent or subarborescent woody bam- 
boos with leptomorphic (running) rhizomes, persistent to deciduous and mostly 
glabrous culm leaves, and leaves at the tip of new shoots crowded into a dis- 
tinctive fan-shaped cluster or top knot with blades expanded as on foliage leaves. 
Branch complements typically have | primary branch and 0-2 subequal sec- 
ondary branches arising from shortened internodes at the base of the primary 
branch, which rebranch to produce up to 40 or more secondary branches on 
older culms. The culm and foliage leaves bear fimbriae and usually also au- 
ricles. Synflorescences in this group are determinate, open, and racemose or 
paniculate, with 6-12 laterally compressed florets per spikelet. Spikelets have 
1-2 glumes and 3 stamens per floret. Like most temperate bamboos, 
Arundinaria has a basic chromosome number of x = 12 and presents several 
enigmatic characteristics including delayed flowering and monocarpy; repro- 
duction is primarily vegetative and seed production is infrequent and unpre- 
dictable. Arundinaria ss. is confined to southeastern portions of the continen- 
tal United States (specifically Alabama, Arkansas, Delaware, Florida, Illinois, 
Indiana, Kentucky, Louisiana, Maryland, Mississippi, Missouri, New Jersey, 
North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia, 
and West Virginia), occurring from the Coastal Plain from New Jersey south to 
Florida and west to eastern Texas, and inland through the Piedmont to moder- 
ate elevations in the Appalachian Mountains. Arundinaria once formed exten- 
sive and dense canebrakes (with or without tree canopy) covering vast areas of 
fertile river bottomland often described by early explorers in the pre-colonial 
U.S. (West 1935), but has been greatly reduced in extent and abundance from 


TRIPLETT ET Al 81 


its historical range by grazing and fire suppression (Hughes 1951, 1957, 1966; 
Platt & Brantley 1997; Judziewicz et al. 1999). 

Two species (Arundo gigantea Walter, Arundo tecta Walter) were originally 
described by Walter (1788) and represent extremes of morphological types 
found among inland and coastal forms, sometimes referred to as “Mississippi- 
type” and “Atlantic-type’ aan 1943), pees eeae ee 
(river cane or giant cane) forms extensive nies in low woods, moist ground, 
and along riverbanks from the lowlands east tof the Appalachians, west to Mis- 
souri, up the Mississippi Valley to southern Illinois and up the Ohio River to 
southern Ohio. Arundinaria tecta (Walt.) Muhl. (switch cane) forms colonies 
in non-alluvial swamps, moist pine barrens, live oak woods and along sandy 
margins of streams, preferring moister sites than A. gigantea. It is restricted to 
the Coastal Plain of the southeastern U'S., from southern Maryland to Alabama 
and Mississippi (McClure 1973; Hitchcock 1951). 

Within Arundinaria s.s., complex population-level variation has contin- 
ued to be problematic for taxonomists and field botanists (McClure 1973; 
Judziewicz et al. 1999; Platt & Brantley 1997). Phenotypic diversity among North 
American cane populations has inspired diverse taxonomic interpretations, 
generally with 1-3 taxa recognized at either specific or subspecific levels (Gilly 
1943; Young & Haun 1961; Voight & Mohlenbrock 1964; Radford et al. 1968; 
Hitchcock 1971; McClure 1973; Campbell 1985; Tucker 1988; Platt & Brantley 
1997). McClure (1973) published the most recent exhaustive treatment of 
Arundinaria and took a conservative approach, recognizing a single polymor- 
phic species (A. gigantea) and three subspecies, one of which [A. gigantea ssp. 
macrosperma (Michx.) McClure] is a catch-all for putative hybrids derived from 
the introgression of the other two. However, McClure acknowledged that fur- 
ther studies, particularly genetic-based studies, were necessary to clarify the 
phylogeny and taxonomy of this group. 

In the Southern Appalachians, astute botanical observers have long ques- 
tioned the identity of a curious short-statured cane that typically occupied sites 
away from streams and rivers. Among the diverse North American populations 
of Arundinaria, a variety with deciduous foliage was recorded by botanist C.D. 
Beadle in Western North Carolina (Beadle 1914; see also Young 1945). Beadle 
recognized this form as A. tecta var. decidua, not ruling out the possibility that 
it might in fact be a distinct species. It is unclear why Beadle associated this 
deciduous variety with A. tecta, although perhaps it was because of the small 
stature typically associated with A. tecta. In the first half of the nineteenth cen- 
tury, two botanists affiliated with the University of North Carolina Herbarium, 
William Willard Ashe and William Chambers Coker, made notes on specimens 
collected near Highlands, North Carolina indicating that the short, delicate, 
deciduous cane of the mountains might be a distinct taxon. Roland Harper 
(1928) was also intrigued by cane specimens occurring on bluffs in northern 


82 BRIT.ORG/SIDA 22(1) 


Alabama and thought that an unrecognized taxon might be present. In the lat- 
ter half of the twentieth century, botanists and ecologists have informally con- 
sidered the small, upland cane as “hill cane” and have been unconvinced that it 
could be assigned to either A. gigantea or A. tecta. Hill cane is often common in 
mesic and submesic Sones and upland woodlands. Moreover, because of over- 
lapping morphological characteristics, floristic descriptions of A. tecta and A. 
gigantea may be confounded by this distinct form of cane. For example, 
Arundinaria tecta has been described as occurring along river branches 450 to 
580 feet above sea-level on the southeast slopes and along the courses of moun- 
tain streams and shady mesic hillsides in the foothill region, well back from 
water (Harper 1928; Peattie 1929), but these inland and upland habitats almost 
certainly are populated by hill cane, not switch cane. In our field work on 
Arundinaria, we located several widespread populations of hill cane along the 
southern Appalachian Mountain chain, extending the range of the form that 
Beadle described. Field work has been complemented by herbarium studies to 
produce our current understanding of the range for this entity. 

Species limits within Arundinaria s.s. have not been examined previously 
ina phylogenetic framework or with molecular tools. Our investigation of hill 
cane is part of a larger study of the phylogenetic history of the North Temper- 
ate bamboo clade (in collaboration with the Bamboo Phylogeny Group), and as 
part of that study we are reconstructing the phylogeny of river cane, switc 
cane, and hill cane utilizing Amplified Fragment Length Polymorphism (AFLP) 
data to test the monophyly of putative species and to correlate the results with 
morphological and geographical characters, with the goal of producing a re- 
vised treatment of the genus. Ordination analyses (PCA) of mot phological char- 
acters and AFLP studies will be presented in a later publication. 

Preliminary cladistic analysis of AFLP data (Triplett & Clark, in prep.) dem- 
onstrated that the three types of cane form separate monophyletic lineages en- 
compassing two previously recognized entities (river cane and switch cane) and 
one entity encompassing those plants recognized as hill cane. Moreover, hill 
cane specimens from a wide geographic range cluster as the sister clade of A. 
tecta, rejecting the hypothesis that hill cane is an ecologically induced form of 
A. tecta and instead suggesting that it is a distinct lineage with a unique evolu- 
tionary history. These results prompted a reevaluation of diagnostic characters 
within Arundinaria s.s. Preliminary ordination (PCA) analyses of morphologi- 
cal characters similarly identified three non-overlapping entities and allowed 
us to recognize the most important diagnostic vegetative features. These groups 
correspond precisely with the three lineages derived from the AFLP data. We 
therefore propose the recognition of each entity at the species level: A. gigantea 
(river cane), A. tecta (switch cane) and a previously undescribed species (hill 
cane). In advance of the publication of the Flora of North America, we here 
describe and illustrate the new species, A. appalachiana Triplett, Weakley, & 


—_ 
=) 


TRIPLETT ET AL., A NEW SPECIES OF 83 


L.G. Clark, from the southern Appalachian Mountains, and compare and con- 
trast it with its congeners A. gigantea and A. tecta. 


MATERIALS AND METHODS 


Field studies of natural populations conducted in October 2003 and July- 
October 2005. Standard bamboo collection procedures were followed 
(Soderstrom & Young 1983); bulky specimens of rhizomes, branch comple- 
ments, and culm nodes and internodes were made for all collections. 

Herbarium specimens from A, F, GA, GH, ISC, NCU, and US (herbarium 
acronyms following Holmgren & Holmgren 2006) were examined. While our 
taxonomic circumscription of Arundinaria s.s. is based on an approach com- 
bining morphological and molecular data, we have relied upon morphological 
characters to provide identifications of the specimens examined. Complete 
specimens, including culm leaves, buds, branch complements with foliage 
leaves, and synflorescences were rarely available, and some herbarium speci- 
mens of Arundinaria could not be conclusively assigned to species. Only one 
flowering specimen was located among the specimens identified as A. 
appalachiana. 

Specimens were measured for a variety of morphological characters, in- 
cluding foliage leaf length and width, inner ligule length, inflorescence length, 
spikelet length, and lengths of spikelet bracts (glumes I-IV, lemma and palea). 
Top knot (the cluster of leaves at the tip of new shoots) and foliage leaf lengths 
were measured from the base of the pseudopetiole to the tip of the blade. Leaf 
width was measured at the widest point. Primary branch length was measured 
from the point of origin at the node to the end of the branch axis. Synflorescence 
length was measured from the base of the basalmost branch to the apex of the 
main axis. Spikelets were removed from specimens and softened using a modi- 
fied Pohl’s solution (Pohl 1965: 750 ml distilled water, 250 ml 1-propanol, 2 ml 
liquid dish soap), dissected, examined, and measured for floral characters us- 
ing a dissecting microscope equipped with a micrometer. Anatomical charac- 
ters of leaf blades (both epidermal micromorphology and cross sections) were 
obtained using light microscopy of sections made following standard proto- 
cols for free hand sectioning and epidermal peels (Clark 1986; Ellis 1976, 1979). 


RESULTS AND DISCUSSION 


Our decision to recognize this taxon at the species level is based upon the com- 
bination of phylogenetic and morphologic analyses with careful consideration 
of the decisions made in the past regarding the North American Arundinaria 
species complex and the ability to diagnose monophyletic units. This interpre- 
tation follows from morphological (ie., diagnostic characters) and phylogenetic 
(ie. unique ancestry) species concepts (Olmstead 1995; Sites & Marshall 2003): 
The features discussed below are those identified as the most diagnostic based 


84 BRIT.ORG/SIDA 22(1 


on observations made during field work and morphological ordination analy- 
ses (PCA). The most consistent differences among the North American species 
are seen in vegetative characters including features of the rhizomes, culm in- 
ternodes and culm leaves, branching, and top knot and foliage blades, described 
below and summarized in Table 1. 


Distribution & ecology 

Arundinaria appalachiana is indigenous to the southern Appalachian Moun- 
tains where it occurs in the southern Blue Ridge, Blue Ridge /Piedmont Escarp- 
ment, upper Piedmont, and Ridge and Valley physiographic provinces (Fig. 1). 
The full extent of its distribution is still poorly known, because of the infre- 
quent collection of bamboos in eastern North America and the of ten poor qual- 
ity of the existing specimens; for this reason we have chosen to supplement 
vouchered specimens with additional county records based on what we con- 
sider reliable sight records of this new species (these counties should be veri- 
fied with vouchers). Hill cane iscommon in oak-hickory forests and woodlands 
on mesic, submesic, and xeric slopes and uplands, sometimes occurring as well 
in hillside seepages, but nearly always on slopes, bluffs, and ridges away from 
perennial streams, in contrast to the geographically sympatric A. gigantea. As- 
sociated overstory species include Quercus montana Willd., 9. coccinea Mtnchh., 
Q. alba L., Q. rubra L., Q. falcata Michx., Pinus echinata Mill. P virginiana Mill, 
Carya alba (L.) K. Koch, Carya glabra (Mill.) Sweet, and Nyssa sylvatica Mar- 
shall. Although these slope and ridge forests are well-drained, the annual rain- 
fall amounts in this region are high and equably distributed. In the escarpment 
gorges of the Chattooga River, Whitewater River, Thompson River, Horsepasture 
River, Toxaway River, Horsepasture River, and Eastatoe Creek, the highest an- 
nual rainfalls in eastern North America (exceeding 80 inches a year) maintain 
higher then expected moisture levels even in topographic situations that tend 
tocreate xeric communities (ridgetops and convex upper slopes and side ridges) 
(Robinson 2000). Still, there is no question that hill cane occupies unusually 
dry and upland habitats compared to its congeners. Arundinaria gigantea typi- 
cally occurs on the floodplains of large to small rivers, sometimes edging onto 
lower portions of mesic slopes, whereas A. tecta typically occurs along small to 
medium blackwater rivers, in swamps, on deep peat in pocosins, and in small 
seepages with organic soils. 

Arundinaria dppalachiana is sympatric (at least in the broadest sense) 
with both of its congeners, in that it occurs in the same counties and within a 
few kilometers of populations of both A. gigantea and A. tecta. Within the re- 
gion occupied by A. appalachiana however, A. gigantea occurs only along the 
upper reaches of major rivers, notably the Little Tennessee and French Broad 
Rivers, while A. tecta also occurs at lower elevations and in different topographic 
and ecological situations. 


TRIPLETT ET AL., A NCW SF EXITED UF 


Taste 1. Morphological comparisons of A 


dinaringn 


palachiana,A. tecta, and A. gigantea. 


Character A. appalachiana A. tecta A. gigantea 
Rhizome air canals present or absent — present absen 
ulcus usually absent usually absent et present 
Culm leaf duration persistent persistent eciduous 
Culm leaf auricles absent present, deciduous present, 
deciduous 
Top knot number of leaves 6-12 9-12 - 
Top knot leaf blade length (cm) 9-22.5 20-30 16-24 
Compressed basal internodes 2-5 2-4 0-1 
n primary branch 
1° branch basal nodes: 2° branches absent present,subequal — present, subequal 
Primary branch length (cm) 7-33 usually >50 15-25 
Foliage leaf blade length (cm) 5-20 7-23 8-15 
Foliage leaf blade width (cm) 8-2 1-2 0.8-1.3 
Foliage leaf vestiture pilose or glabrous densely pubescent densely pubescent 
or glabrous or glabrous 
Foliage leaf duration deciduous evergreen evergreen 
Foliage leaf texture chartaceous coriaceous subcoriaceous 


Foliage leaf abaxial tessellation 


weakly tessellate 


strongly tessellate 


| 1] 
DUUT ly LESSEN GLO 


The species biology of A. appalachiana is poorly understood. Its congeners 
are long-lived monocarpic perennials, and this appears to be the case with A. 
appalachiana as well. It has been seen flowering and fruiting even more infre- 
quently than its congeners; judging from its habitat, its flowering and fruiting 
may be stimulated by fire (like A. tecta), and the paucity of fertile specimens 
may reflect the general suppression of fire in its habitat through the twentieth 
century. Field observations suggest that clones of A. appalachiana are slow- 
growing and very long-lived, certainly persisting for decades and likely for cen- 
turies. 


Morphology 
Rhizomes.—The rhizomes of A. appalachiana are leptomorphic, a characteristic 
they share with other north temperate woody bamboos; however, in many cases 
the growing tips of new rhizomes travel only a short distance before turning up 
to form a new culm, thus presenting a sympodial branching pattern. This pat- 
tern also occurs in A. tecta but has not been confirmed for A. gigantea. An inter- 
esting characteristic of this species is the variability in air canal development 
(McClure 1963). Air canals are present in some specimens but not others, and 
may in fact be longitudinally and peripherally discontinuous in rhizomes of A. 
appalachiana. Air canals are ntly 
continuous. 

Culm internodes.—Although the culm internodes of A. appalachiana can 


lyf present in A. tecta and 


desta Be 


86 BRIT.ORG/SIDA 22(1) 


Ree Py ae 


aie. ee 
Ceae ar co aewen aaa 
Syste ea NI Oy 
anes serene 

Ee to ee 
sta oo KD 
RES 


be somewhat flattened behind the branch complement, the internodes lack a 
prominent groove or sulcus. This is consistent with A. tecta but contrasts with 
A. gigantea, which typically has internodes that are prominently sulcate. 
Branching.—In bamboos, the morphology and architecture of the set of 
branches arising from culm nodes (the branch complement) is a source of nu- 
merous taxonomically useful characters. In Arundinaria, the pattern of short- 
ened or compressed internodes at the base of primary branches and the extent 
and pattern of secondary branching are especially valuable. The branch comple- 
ment of A. appalachiana is characterized by 2-5 shortened or compressed in- 
ternodes at the base of the primary branch, without rebranching in this basal 
area. The first elongated internode above the shortened ones is typically con- 
strained to ~30% the length of distal internodes. In contrast, A. tecta, while 
having a similar pattern of compressed internodes, typically will produce buds 
and branches from the nodes in the area of compression, creating subequal 
branches from the base of the primary branch. Arundinaria gigantea typically 
has only one (or no) compressed basal internode, but if present, this node may 
produce a secondary branch. Primary branches in A. appalachiana are usually 


TRIPLETT ET Al 87 


less than 35 cm long. In contrast, A. tecta produces long primary branches usu- 
ally >50 cm. 

Culm leaves.—The culm leaves of A.appalachiana are typically shorter than 
their associated internodes at the base of the plant, becoming progressively 
longer towards the top knot. At midculm they are approximately the same 
length as the associated internode. In contrast, midculm culm leaves of A. tecta 
are longer than their associated internodes, and those of A. gigantea shorter. 
Arundinaria appalachiana and A. tecta have persistent culm leaf sheaths, 
whereas A. gigantea has deciduous sheaths. The culm leaf sheaths of A. 
appalachiana are tessellate; however, their tessellation is not as pronounced as 
itisin A. tecta. The culm leaves lack well-developed, prominent auricles, unlike 
A. gigantea and A. tecta. 

Top knot and foliage leaves.—In Arundinaria, leaves at the tip of new culms 
are crowded into a distinctive fan-shaped cluster or top knot, with their blades 
expanded as on foliage leaves. The top knot leaf blades of A. appalachiana are 
typically 9-22.5 cm in length, while A. tecta typically has larger blades (20-30 
cm long); those of A. gigantea are typically 16-24 cm in length. The foliage leaf 
blades of A.appalachiana are deciduous; bladeless branches persist on the plants 
after leaf drop, often with the older sheaths still intact. The blades are 
chartaceous, presumably, since the blades are deciduous, the plant invests less 
energy in producing them by producing less sclerenchyma. In contrast, the 
leaves of A. gigantea are persistent and subcoriaceous, while leaves of A. tecta 
are persistent and coriaceous. The abaxial surfaces of the leaf blades of A. 
appalachiana are weakly tessellate, whereas in A. gigantea and A. tecta they are 
strongly tessellate. The abaxial and adaxial leaf surfaces are typically sparsely 
to more or less densely pilose (glabrous in some populations) in A. appalachiana. 
Leaf blades of A. gigantea are typically densely pubescent with short, soft hairs 
on abaxial surfaces, while the blades of A. tecta are densely pubescent on both 
surfaces; neither of these pubescence patterns has been seen in A.appalachiana 

It is important to note that Arundinaria gigantea and Arundinaria tecta 
can both survive in deeper shade in the forest, albeit in stunted conditions, and 
thus it is possible to find diminutive or depauperate plants of both that super- 
ficially resemble hill cane, mostly in stature. However, a combination of char- 
acters such as the branch complement, leaf texture, and leaf pubescence are 
usually sufficient to distinguish these stunted forms of river and switch cane 
from hill cane populations and from each other 


TAXONOMIC TREATMENT 
KEY TO THE SPECIES OF ARUNDINARIA SENSU STRICTO 


1. Primary branches with 0-1 compressed basal internodes; culm internodes usually 
sulcate; culm leaves deciduous A. gigantea 


88 BRIT.ORG/SIDA 22(1 


1. Primary branches with 2-5 compressed basal internodes; culm internodes usually 
terete; culm ies Salsa to oy cele UOUS. 
2. Foliage bl Is, persiste f ly pubescent or gla- 
brous, elk tessellate; tae ae eae usually more than 50 cm long, basal 
nodes developing secondary branches; top sus blades a. 30. cm long A.tecta 
2: ieee we hartaceous, deciduous, al | glabrot Is. Weakly 
branches usually less than 35cm ong. basal ode: not devel- 
oping secondary branches; top knot blades 9-22.5 cm long A. appalachiana 


Arundinaria appalachiana Triplett, Weakley @ L.G. Clark, sp. nov. (Figs. 1-5). 
TYPE: Ue STATES. ALABAMA: Dekalb Co.: Desoto State Park by Rt. 89 (34° 30' N Lat; 85° 
38' W Long.), ~ ca. 515 m, 25 Jul 2005, Triplett & Ozaki 99 (HOLOTYPE: ISC; ISOTYPES: MO, 


NCU, UNA, U 
Rhizomata leptomorpha, strat l eri i lum praesenti.Culmi 2-6 mm diametro, 
0.5-1.8 m alti, omnino glabri, erecti. nternodia 4.5-12 cm ronga te teretia ailgine sulco). Vaginae culmorum 
5.5-11(-15) cm longae, persistentes, sine auriculis. Culmi j inali(5-)6-12 foliorum, 


laminis (5-)9-22.5 cm longibus, 1.4-2.8 cm latibus, linear bee vel lineari-lanceolatis vel ovati- 
lanceolatis, pilosis vel interdum glabris. Rami pew ad nodos medianos c I 3 en 
longi, internodiis abbreviatis sine gemmis basi 2-5. Folia eae complementi 3-7. Laminae 
foliorum (3-)5-20 cm longae, 0.5-2 cm latae, shanties ae, deciduae, pilosae vel interdum glabrae, 
abaxialiter infirme tessellatae. Synflorescentiae 7-11.5 cm ee 2-5 cm latae, determinatae 
racemosae apertae; pedicelli 4-25 mm longi, spiculis 6-8. Spiculae 3-5.5 cm longae, glumis 1(-2), 
flosculis basilibus interdum sterilibus, 5-8 flosculis fertilibus continentibus et anthoeciis 
rudimentalibus terminalibus 1-3; glumae inaequales, 5-nervatae, attenuatae, glabrae; gluma I 3-6.5 
mm longa; gluma II 5.5-9 mm longa; lemmata fertilia 11-16 mm longae, 7-1l-nervata, apicibus acutis 
vel acuminatis, abaxialiter glabris; palea LO-13 mm longa, 8-10-nervata. Stamina 3; anthera 5-7 mm 


Woody bamboo. Plants of diffuse to (pluri-) caespitose habit. Rhizomes 
leptomorphic, usually horizontal for a only short distance before turning up at 
the apex to form a culm (therefore often presenting a sympodial branching 
pattern), hollow (with a small central lumen), peripheral air canals sometimes 
present (if so, apparently discontinuous longitudinally and/or peripherally). 
Culms 2-6 mm in diameter, 0.4-L8 m tall, erect, tillering; internodes 4.5-12 cm 
long (progressively shorter towards culm apex), terete, hollow, glabrous, flat- 
tened behind the branch complement on larger culms but the sulcus not promi- 
nent; nodes solitary, the nodal line horizontal, supranodal ridge not aoe 
bud one per node (single) on a slight promontory, triangular, the shoulders of 

the prophyll ciliate. Culm leaves persistent, approximately equaling associated 
internodes at midculm, typically shorter than associated internodes at the culm 
base, becoming proportionally longer towards the culm apex; sheaths 5.5-11 
(-15) cm long, shortest on lower nodes, becoming progressively longer towards 
the culm apex, glabrous, margins ciliate; blades 0.5-1.4 cm long, triangular to 
lanceolate, reflexed to erect, glabrous, deciduous, intergrading into top knot 
leaves; auricles absent; fimbriae 1-9 mm long, ascending to erect; inner ligules 
ca. 0.1 mm long, a fringe of short cilia; outer ligule absent. Top knot leaves in an 
apical cluster of (5-)6-12; sheaths glabrous, margins ciliate; auricles absent; fim- 


90 BRIT.ORG/SIDA 22(1) 


Fic. 3. Arundi hiana. A. Partially di I florets. B. Synfl with five spike- 
lets. Scale bar = 1 cm. m. Drawings based on Ahles & Leisner 15147. (Illustrations by J. Triplett) 


briae 1-9 mm, ascending to erect; blades (5-)9-22.5 cm long, 1.4-2.8 cm wide, 
L:W = 8.3-9.3, linear, linear-lanceolate or ovate-lanceolate, chartaceous, pilose 
or glabrous, abaxially weakly tessellate, apices acuminate, bases attenuate to 
cuneate, midrib + centric. Branching infravaginal (rarely extravaginal); primary 
branches | per node, 7-33cm long, with 2-5 compressed basal internodes, basal 
nodes not developing secondary branches; first elongated internode shorter than 
subsequent ones (~30%); higher order branches present on older plants, reiter- 
ating the 1° branch (ie., with the same pattern of compressed basal internodes 
and branching). Foliage leaves 3-7 per complement; sheaths glabrous, margins 
ciliate, weakly tessellate; auricles absent; fimbriae 1-9 mm, ascending to erect; 
inner ligule glabrous or ciliate, fimbriate or lacerate; outer ligule present as a 
minute rim: blades linear, linear-lanceolate, or ovate-lanceolate, chartaceous, 
deciduous, surfaces pilose (sometimes glabrous), abaxially weakly tessellate, 
apices acuminate, bases attenuate to cuneate, midrib + centric; primary branch 
foliage leaf blades (4-)9-20 cm long, (0.5-)0.8-2 cm wide; L:W =10.7-1L7; higher 
order branch foliage leaf blades (3-)5-17.5 cm long, (0.5-)0.8-15 cm wide; ter- 
minal foliage leaf blade aaa unexpanded laterally, pasa: but persisting 
as a tail-like appendag 7-11.5cm long, 2-5 cm wide, determi- 


wo 


TRIPLETT ET AL., ANEW SPFEAILS UP 91 


Fics. 4-5. Arundinaria appalachiana. 4. Habit, in Rhea Co., Tennessee. 5. Close-up of primary branch with compressed 
basal internodes, Dekalb Co., Alabama. (Photos by J. Triplett) 


nate, open-racemose, apparently borne on specialized nonleafy shoots; peduncle 
10-17 cm long (3 measured), glabrous, terete; rachis glabrous; pedicels 4-25 mm 
long; 6-8 spikelets per synflorescence. Spikelets 3-5.5 cm long, laterally com- 
pressed, disarticulating above the glumes and between the florets, consisting 
of 1(-2) glumes, occasionally a basal sterile floret, 5-8 fertile florets and 1-3 
progressively rudimentary apical sterile florets; rachilla internodes 3-4 mm 
long; glumes unequal, 5-nerved, attenuate, glabrous; glume I 3-6.5 mm long; 
glume II 5.5-9 mm long; fertile lemmas 11-16 mm long, 7-1l-nerved, apex acute 
or acuminate, abaxially glabrous, transverse veinlets barely perceptible or not 
at all manifest, usually somewhat reddish-purple; paleas 10-13 mm long, 8-10- 
nerved, broadly sulcate and 2-keeled dorsally; lodicules and ovary not seen. Sta- 
mens 3; anthers 5-7 mm long. Fruit not seen. 

Distribution and Ecology.—(Fig. 1). Endemic to the southern Appalachians 
and upper Piedmont of northeastern Alabama, northern Georgia, southwest- 
ern North Carolina, northwestern South Carolina, and southeastern Tennessee, 
United States; 300-800(-1065) m. In upland oak-hickory-pine forests on slopes, 
less typically in more mesic sites, seeps, or along small streams. 

Phenology.—Lack of specimens in flower or information on the extent of 


92 BRIT.ORG/SIDA 22(1) 


blooming makes it impossible to determine flowering behavior in this species 
at present. Of the specimens cited below, only one confirmed flowering speci- 
men of this species was identified, suggesting that flowering may be an even 
rarer event in this species than in most woody bamboos. 
Etymology.—Arundinaria appalachiana is named for its distribution in the 
forests of the Appalachian Mountains. 
Common name.—hill cane. 


R ined: UNITE D STATES, ALABAMA. Dekalb Co.: Little River Canyon 
National peste plot PIEC 27, 2002, McDaniel s.n.(NCU); in steep wooded bank of West Fork, Little 
River, across stream from overlook shelter, DeSoto State neh 9 May 1959, Sherman & Carter 25747 
(GH); Desoto State Park on trail by Laurel Creek (34° 30'N Lat; 85° 38’ W Long.), elev. ca. 515m, 25 Jul 
2005, Triplett & Ozaki 100 USC, MO, US). GEORGIA. Bartow Co.: in woods on S side of Stamp Creek 
Rd. just E of Jones Mill Rd. (34° 14' N Lat; 84° 40' W Long.), elev. ca. 311 m, 26 Sep 2005, Triplett 166 
CISC, MO, US). Rabun Co.: maple-birch-magnolia association, Laurel Creek Olivine Deposit, 1.5 mi E 
of Pine Mt., 21 Jun 1946, Radford s.n. (NCU ): + -hickory woods on Pine Mt., Bald Rd., 4 Jun 1952, 
Radford 6134 (NCU); Warwoman Road and Overflow Creek bridge, N side, 20 May 1996, Stancil 950 
(GA); in swamp near top of Oakey Mountain, SE of Nacoochee Reservoir, 25 May 1941, Duncan 3283 
(GA); on trail from 155 0.5 mi S of Kattie Place, elev. ca. 790 m, 22 Oct 1995, Milsted & Zhang 585 (GA); 
pine-oak woodland near roadside park, | mi N of Tallulah Falls River, 9 May 1967, Isely, Welsh, & Isely 
10286 USC), 2 mi N of Warwoman Rd. along GA 28, scattered aan mature woods W of road 
(34° 57'N Lat; 83° LO’ W Long.), elev. ca. 687 m, 25 Oct 2003, Triplett & Clark 20 (SC, MO, US). Stephens 
Co.: Deep gorge, Cedar Creek, Camp Mikell Rd. off GA 184, just W of Camp Mikell and N of Toccoa 
(34 08' 06" N. lat; 83 20' 13” W. long), elev. ca. 300 m, 19 Jun 1975, Boufford & Wood 16501 (NCU), NW- 
facing ravines and ridges on the S side of Panther Creek, SW of Yonah Lake (Tugaloo River), N of 
Toccoa (34 43' 30” N. lat; 83 21' 13” W. long.), 25 Jun 1975, Boufford & Wood 16766 (NCU). NORTH 
CAROLINA. Clay Co.: Oak-hickory woods, 4 mi NW of Hayesville, 5Jun 1952, Radford & Wood 6162 
(NCU). Gaston Co.: low woodland near the northern tip of Pasour Mt., about 3 mi SSW of High Shoals, 
21 Jun 1956, Ahles & Leisner 15147 (NCU); oak-hickory woods on Crowder’s Mt, 4 Jun 1953, Radford 
7084 (NCU). Graham Co.: small population on N side of Santeetlah Dam Rd. (SR 1146) just off Hwy 
129 (35° 22'N Lat; 83° 5I’ W Long.), elev. ca. 537 m, 2 Oct aie eran MO, US). Jackson Co. 
on highway between Dillsboro & Park Entranc n e top of bank by roadside, 26 


= 


Jul 1937, Coker s.n. (NCU); Cope Creek ae ne af NC 23 out ci ae growing among Polytrichum 

sp., Pine, and sedge (35° 23' N Lat; 83° Long.), elev. ca. 675 m, 25 Oct 2003, Triplett & Clark 21 
CISC, MO, US). Lincoln Co.: rich oe forest and stream banks 0.4 mi W of Cat Square, 28 Apr 
1957, Bell 6638 (NCU). Macon Co.: maple-birch-magnolia (cove) association, Corundum Hill Olivine 
Deposit, 1 1/2 mi NW Gneiss, 15 Jun 1946, Radford s.n.(NCU); Horse Cove near Highlands, 3 Sep 1948, 
Radford s.n. (NCU); in deep, shaded ravine at top of pasture at foot of Whiterock Mountain, 7 mi from 
Otto, NC on Tessentee Creek Rd., elev. ca. 1065 m, 7 Aug 1938, Stewart & Hechenbleikner s.n. (NCU) 
pine-broom-straw association, Corundum Hill Olivine Deposit, 1.5 mi NW Gneiss, 15 Aug 1946, Rad- 
ford sn. (NCU); Mulberry Rd. ca. 0.5 mi off 441, on steep hillside among oak, rhododendron, and 
maple (35° OLN Lat; 83° 23' W Long.), elev. ca. 647 m, 25 Oct 2003, Triplett, Clark, & Weakley 19 (SC, 


MC . McDowell Co.: near Marion, 25 Oct 1915, Ashe s.n. (NCU). Polk Co.: 4 mi W of Tyron, valley 
of Fall Creek in wet mez dows on eee in reservoir, 19 Jun 1942, Walker 3469 (US). Rutherford 
Co.: wet ditch on CR 1721 2 mi N of Sunshine, 24 Jun 1967, Smith 74 (NCU). Swain Co.: Bryson City, 


by stream, 1] Jul 1927, Hunne a sn. (GH); Bryson City, private property on W side of Wiggins Rd., 
0.3 mi up from Betts Branch (SR 1343) (35° 26' N Lat; 83° 25' W Long.), elev. ca. 601 m, | Oct 2005, 
Triplett 184 (SC, MO, US). Transylvania Co.: rocks and cliffs on side of mountain, 100 to 200 feet 
ee the bank of Davidson River, Pisgah Forest, 28 Sep 1915, Ashe s.n. (NCU); Pisgah Forest, roac 


to 


TRIPLETT ET AL., A 93 


Pink Beds, 23 Aug 1938, Stewart s.n. (NCU); Middle Bearcamp Creek area, roadside, Highlands, 2 Aug 
1962, Rodgers & Shake 62165a (NCU), oak woods near Looking Glass Falls, 7 Jun 1952, Radford & 
Wood 6192 (NCU); Horsepasture Gorge, roadside about 1 mi W of crossing, elev. ca. 488 m, oyun 1961, 
Rodgers 6162a (NCU). SOUTH CAROLINA. Greenville Co.: Cedar Mountain, 0.4 mid road 

mi S of Caesar's Head SP Visitor Center (35° 05' N Lat; 82° 36’ W Long,), ae ca.515 m, 30 Sep 2005, 
Triplett 179 (SC, MO, US). Oconee Co.: oak-hickory woods ridge above Walhalla Fish Hatchery, ca. 11 
mi N of junction of S.C. Routes 28 and 107 on Route 107, 9 Jun 1952, Wood 7879(A); Hill property, NW 
side Old Rocky Gap Road, W side West Village Creek near creek, Mountain Rest, Blue Ridge prov- 
ince, elev. ca. 520 m., 24 Aug 1991, Hill 22585 (GH). Pickens Co.: mixed deciduous forest, 3 mi N of 
pe Bottom near US 178, 22 Aug 1956, ta 16758 (NCU); Boggs’ Rock, granite-gneiss outcrop N 

of Liberty, 3 Jun 1974, Knox 407 (NCU) wooded slope, 2.4 mi S of NC line on US 178, 8 Jun 1956, 


small plant at summit of mountain, in hardwood forest. 29 Jun 1957, Pohl 7664A (ISC). Monroe Co.: 
White Cliff Springs, Jul 1890, Lamson-Scribner s.n. (US). Polk Co.: Boyd Gap Overlook above Ocoee 
River, 4.2 mi W of Hwy 68, on steep wooded hillside just beyond entrance to Boyd Gap Trail #331 
(35° 02' N Lat; 84° 27' W Long.), elev. ca. 529 m, 26 Sep 2005, ae 165 (SC, MO, US). Rhea ve 
Firetower Rd just off Hwy 68 next to Grandview Community Center and below intersection wit 
Emergency Road (35° 44’ N Lat; 84° 50’ W Long.), elev. ca. 445 m, 3 Oct ee Triplett 188 (ISC, MO, 


ACKNOWLEDGMENTS 


Field work was supported by a Graduate Research Grant from the American 
Society of Plant Taxonomists and grants from the American Bamboo Society to 
JT as well as a National Geographic Society Grant (#7336-02) to LC. Support 
for JT and for the final preparation of this manuscript came from a National 
Science Foundation Grant (DEB-0515712) to LC. We are indebted to Angus 
Gholson (Chattahoochee, FL), Michael Hotchkiss (USDA, Byron, GA), Robert 
and Lauren Clark (Peachtree City, GA), Jim and Frankie Triplett (Winston-Sa- 
lem, NC), Gary Kauffman, Linda Lee Leslie, Adam and Sue Turtle (Earth Advo- 
cates Research Farm, Summertown, TN), and Julian Campbell (TNC, Lexing- 
ton, KY) for their hospitality or assistance in locating populations of 
Arundinaria. We also thank the curators of the herbaria that loaned specimens 
for this study. We are grateful to Gary Kauffman, Keith Langdon, Virginia 
McDowell, Tom Govus, and Mike Schafale for their interest and assistance in 
resolving the persistent taxonomic issue of the correct identity of the small 
upland mountain cane. Chris Stapleton (Royal Botanic Gardens, Kew) kindly 
pointed out to us the importance of compressed internodes in the taxonomy of 
this group and Emmet Judziewicz (Dept. of Biology, University of Wisconsin- 
Stevens Point) suggested the specific epithet. 


REFERENCES 


Beaote, C.D. 1914. Bamboo. In: Bailey, L.H. The standard cyclopedia of horticulture, Vol. 1. 
New York: The Macmillan Company. P. 446. 

Campsett, J.N. 1985. The land of cane and clover: presettlement vegetation in the so-called 
bluegrass region of Kentucky. Herbarium Report, University of Kentucky, Lexington. 


94 BRIT.ORG/SIDA 22(1 


Car, L.G. 1986. Systematics of Chusquea Section Chusquea, Section Swallenochloa, Sec- 
tion Verticillatae, Section Serpentes, and Section Longifoliae (Poaceae: Bambusoideae). 
Unpublished Ph.D. dissertation, lowa State University, Ames. 

CiarK, L.G.,S. DRANSFIELD, J. TRIPLETT, aNd J.G. SANCHEZ- ey (In eiese): Phylogenetic relationships 
among the one-flowered, determinat f Bambuseae (Poaceae: Bambusoideae). 
In:Columbus, J.T.,E.A. Friar, C.W. rion JM. es L.M.Prince,and M.G.Simpson,eds 
Monocots: comparative biology and evolution. 2 vols. Rancho Santa Ana Botanic Gar- 
den, Claremont, California, U.S.A. 

Eiuis, RP. 1976. A procedure for standardizing comparative leaf blade anatomy in the 
Poaceae. |. The leaf blade as viewed in transverse section. Bothalia 12:65-109. 

Eis, R.P. 1979. A procedure for standardizing comparative leaf blade anatomy in the 
Poaceae. Il. The epidermis as seen in surface view. Bothalia 12:641—-672 

Gity, C.L. 1943. A preliminary investigation of the North American canes (Arundinaria). 
Bull. Torrey Bot. Club, 70:297-309. 

Guo, Q.and R.E. Rickters. 2000. Species richness in plant genera disjunct between temper- 
ate eastern Asia and North America. Bot.J. Linn. Soc. 134:401- 

Harper, R.M. 1928. Economic botany of Alabama.Part 2.Catalogue of the trees, shrubs, and 
vines of Alabama, with their economic properties and local distribution. Geol. Survey 
Alabama Monogr. 9:1-357. 

HitcHcock, A.S. 1951. Manual of grasses of the United States. Rev. by Agnes Chase. 2nd ed. 
USDA Misc. Pub. 200, U.S. Government Printing Office, Washington, DC. 

HitcHcock, A.S. 1971. Manual of the grasses of the United States. Dover Publ., New York. 

Hoimaren, PK.and N.H.Houmeren. 1998 onwards (continuously updated). Index herbariorum. 
New York Botanical Garden. http://sciweb.nybg.org/science2/IndexHerbariorum.asp 

Hucues, R.H. 1951. Observations of cane (Arundinaria) flowers, seed, and seedlings in the 
North Carolina coastal plain. Bull. Torrey Bot. Club 78:113-121. 

Hucues, RH. 1957.Response of cane to burning in the North Carolina coastal plain. North 
Carolina Agric. Exp. Stat. Bull. 402. 

Hucues, RH. 1966. Fire ecology of canebrakes. Proceedings of the Fifth Annual Tall Tim- 

bers Fire Ecology Conference, March 24-25, 1966. Mimeographed. 

uoziewicz, E.J., L.G. Ciark, X. LonDono, and MJ. Stern. 1999. American bamboos. Smithsonian 

Institute Press, Washington, DC. 

KeLCHNeR, S.A.and L.G.Cark. 1997.Molecular evolution and phylogenetic utility of the chlo- 
roplast rp/16 intron in Chusquea and the Bambuscideae (Poaceae). Molec. Phylogen. 
Evol. 8:385-397. 

Li,D.Z.1997.The flora of China Bambusoideae Project: Problems and current understand- 
ing of bamboo taxonomy in China. In: Chapman, G.P., ed. The bamboos. Linnean Soci- 
ety of London Symposium Series. Academic Press, London. 

McCture, F.A. 1963. A new feature in bamboo rhizome anatomy. Rhodora 65:134-136. 
McCiure, F.A. 1973. Genera of bamboos native to the New World (Gramineae: 
Bambusoideae). Smithsonian Contr. Bot. 9. Smithsonian Institution, Washington, D.C 
Oumstead, R.G. 1995. Species concepts and plesiomorphic species. Syst. Bot. 20:623-630. 


TRIPLETT ET AL 95 


Onrneercer, D. 1999. The bamboos of the World. Elsevier. 

Peattie, D.C. 1929. Flora of the Tyron region. J. Elisha Mitchell Sci. Soc 44:141-229. 

Piatt, S.G. and C.G. Brantiey. 1997. Canebreaks: An ecological and historical perspective. 
Castanea 62:8-21. 

Pout, R.W. 1965. Dissecting equipment and materials for the study of minute plant 
structures.Rhodora 67: 95-96. 

Raproro, A.E., H.E. AHLes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill. 

Rosinson, PJ. 2000.Weather and climate. In:Orr, D.M., Jr.and A.W. Stuart. The North Carolina 
atlas: portrait for a new century. University of North Carolina Press, Chapel Hill. Pp. 
21-30. 

Sites, J.W. Jr. and J.C. MarsHALL. 2003. Delimiting species: a renaissance issue in systematic 
botany. Trends Ecol. Evol. 18:462-470. 

SoperstroM, T.R. and S.M. Younc. 1983. A guide to collecting bamboos. Ann. Missouri Bot. 
Gard. 70:128-136. 

Tucker, G.C. 1988. The genera of Bambusoideae (Gramineae) in the southeastern United 
States. J.Arnold Arbor. 69:239-273. 

VoiGHT, J.W. and R.H. MoHLensrock. 1964. Plant communities in southern Illinois. Southern 
illinois University Press, Carbondale. 

Wacter, T. 1788. Flora Caroliniana.J. Fraser, London. 

Wen, J. 1999. Evolution of eastern Asian and eastern North American disjunct distribu- 
tions in flowering plants. Ann. Rev. Ecol. Syst. 30:421-455. 

West, EM. 1935.Canebrakes of the southeastern United States. The Ohio State University 
Press. Abstracts of Doctors’ Dissertations, number 16:253-265. 

Youna, R.A. 1945.Bamboos for American horticulture (I). Natl. Hort. Mag. 24:195-196. 

Youne, R.A. and J.R. Haun. 1961. Bamboo in the United States: descriptions, culture, and 
utilization. United States Dept. Agric., Crops Res. Div., Agric. Handbook No. 193, Wash- 
ington, D.C. 

ZHANG, W.P. and L.G. Ciark. 2000. Phylogeny and classification of the Bambusoideae 
(Poaceae). In: Jacobs, $.W.L.,and J. Everetts, eds. Grasses: systematics and evolution. Pp. 
35-42. CSIRO, Melbourne 


96 BRIT.ORG/SIDA 22(1) 


ANNOUNCEMENT 
The 2006 Applications Delzie Demaree Travel Award 


Applications for the 2006 Delzie Demaree Travel Award should include a letter 
from the applicant telling how symposium attendance will benefit his/her 
graduate work and letter of recommendation sent by the major professor. Please 
send letters of application to: Dr Donna M.E. Ware, P.O. Box 8795, Herbarium, 
Biology Department, The College of William and Mary, Williamsburg, VA 23185- 
8795, U.S.A. 1-757-221-2799, Email: ddmware@wm.edu. The period for receiv- 
ing applications will end three weeks prior to the date of the symposium if a 
sufficient number of applications are in hand at that time. Anyone wishing to 
apply after that date should inquire whether applications are still being ac- 
cepted before applying. The Systematics Symposium dates for 2006 are 13-14 
October. 

The Delzie Demaree Travel Award was established in 1988 honoring Delzie 
Demaree who attended 35 out of a possible 36 symposia before he died in 1987. 
Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His 
teaching career as a botanist began in Arkansas at Hendrix College in 1922. He 
also taught botany at the University of Arkansas, Navajo Indian School, Yale 
School of Forestry, Arkansas AQM, and Arkansas State University at Jonesboro 
where in retired as professor emeritus in 1953. One of the things he enjoyed most 
as a botanist was assisting students with their field botany research. 


— 


SIDA 22(1): 96. 2006 


REVISION OF BAUHINIA SUBGENUS BAUHINIA SECTION 
AMARIA (CERCIDEAE: CAESALPINIOIDEAE: FABACEAE) 


Richard PR Wunderlin 


Department of Biolog 
University of South Florida 
Tampa, Florida 33620-5200 U.S.A 
rwunder@cas.usfedu 


ABSTRACT 


£R 1 ere | R . A 


A taxonomic treatment 
are recognized in series Decandrae andl one species is recognized in series Stenanthae. Bauhinia 
amatlana Wunderlin from southern Mexico, Bauhinia arborea Wunderlin from Ecuador, Bauhinia 


Fifteen species 


ayabacensis wie from Ecuador and Peru, and Bauhinia geniculata Wunderlin from Colombia 
are described and the combination Bauhinia aca eae ‘ Triana ex Hooker f. var. caudigera (S.F 
Blake) Wunderlin is proposed. Keys, descriptions data are provided. 


RESUMEN 


Se presenta un tratamiento taxonomico de Bauhinia subgénero Bauhinia secci6n Amaria. Se 


reconocen quince especies en la serie Decandrae y unaen oe serie ae nae Se edescPen Baubiiie 
d 


amatlana Wunderlin del Sur de México, Bauhinia arborea 
Wunderlin de Ecuador y Pert, y Bauhinia geniculata Wunderlin de Colombia, y se propone la 
combinacion Bauhinia petiolata (DC.) Triana ex Hooker f. var. caudigera (S.F Blake) Wunderlin. Se 
aportan claves, descripciones, y datos sobre la distribucion. 


The pantropical genus Bauhinia with approximately 300 species consists of 
four subgenera: Bauhinia, Piliostigma [=Elayunal, Barklya, and Phanera 
(Wunderlin et al. 1987). Bauhinia subgenus Bauhinia, as defined by Wunderlin 
et al., contains nine sections, of which three (Bauhinia, Pauletia, and Amaria) 
are New Wor 

This is the first of several proposed taxonomic treatments of the New World 
species of Bauhinia. Previous regional treatments in the last half-century such 
as those for Argentina (Fortunato 1986, 1997), Nicaragua (Wunderlin 2001), 
Panama (Wunderlin 1976), western Mexico (McVaugh 1987), Venezuelan 
Guayana (Wunderlin 1998), Middle America (Wunderlin 1983), and parts of the 
genus for Brazil (Vaz 1979; Vaz and Tozzi 2003) will be placed in the larger con- 
text of the entire genus for the Americas following Wunderlin et al. (1987). 

In the following revision, 16 species are recognized as comprising section 
Amaria. Fifteen of these are in series Decandrae and one in series Stenanthae. 
Series Triandrae, containing the single species Bauhinia pansamalana Donn. 
Sm. and placed in section Amaria by Wunderlin et al. (1987), is here transferred 
to section Bauhinia and combined in series Remotae with B. pinheiroi 


SIDA 22(1): 97- 122. 2006 


= 


98 BRIT.ORG/SIDA 22(1) 
Wunderlin with which it shares the characters of three fertile stamens and the 
same pollen type. The pollen sexine of B. pansamalanda is striate-reticulate and 
lacks the infratectal processes characteristic of section Amaria. 


Bauhinia section Amaria (S. Mutis ex Caldas) Endl. Gen. PI. 1318. 1840. Amarias. 
Mutis ex Caldas, Semanario Nueva Granada 2:25. 1810. TYPE 
DC. [=Bauhinia petiolata (S. Mutis ex DC.) Triana ex Hook. | 
ton and Killip (Ann. New York Acad. Sci. 36:164. 1936). 


: Amaria petiolata S. Mutis ex 
|. Lectotype designated by Brit- 


Trees or shrubs; branches lacking infrastipular spines. Leaves unlobed, bilobed, 
or rarely bifoliolate. Inflorescence terminal or axillary; floral bud ellipsoid to 
clavate. Hypanthium cupulate or turbinate to short-tubular; calyx spathaceous, 
sometimes irregularly splitting into several lobes in late aawhiesis petals 5, 
subequal; fertile stamens 10, the inner 5 shorter than the outer 5, or the inner 5 
staminodial, the filaments connate at the base or free; pollen spheroidal, 
inaperturate or 3-colporoidate, the tectum reticulate or rarely striate-reticulate, 
with 


eT 


slunt spine-like or rarely low tectal processes continuous from the foot 
layer through the tectum (infratectal) or rarely supratectal; gynophore partly 
adnate to the abaxial wall of the hypanthium; stigma oblique-capitate. Fruit 
dehiscent. 


KEY TO THE SERIES OF BAUHINIA SECTION AMARIA 


1. Fertile stamens 10; petals spreading 
| 


oO. 


| Series Decandrae 
. Fertile stamens 5; petals strongly imbricate, the corolla forming a pseudotube 


|. Series Stenanthae 


|. Bauhinia series Decandrae Wunderlin, K.Larsen & S.S.Larsen, Biol. Skr. 28:14. 
1987. TypF: Bauhinia petiolata (S. Mutis ex DC.) Triana ex Hook. f 


KEY TO BAUHINIA SPECIES IN SERIES DECANDRAE 
1, Leaves unlobed (rarely bilobate or bifoliolate in 8. beguinoti 
2. Innermost pair of lateral nerves closer to the mid-nerve than to the adjacent 
lateral nerves 
2. Innermost pair of lat 
than to the mid-n 
3. Petals 3.5-4.5 cm Mae fruit 21-27 cm long 
3. Petals 1-2 cm long; fruit 8-13 cm Pu 
4, Leaves 12-30 cm long,6-13 cm wi inal s| lal 
tomentose; fruit elliptic-oblong, 3.5-4 cm wide 
4, Leaves 7-11 cm long, 3-5.5 cm wide; staminal sheath pilose; gynoecium 
glabrous or glandular; fruit linear, 1.5-2.5 cm wide 
1. Leaves bilobate (rarely bifoliolate or unlobed in 8. beguinotil). 
5. Leaves with the innermost pair of lateral nerves closer to the mid-nerve than to 
the adjacent lateral nerves B. beguinotii 
5. Leaves with the innermost pair of lateral nerves equidistant or closer to the ad- 
jacent lateral nerves than to the mid-nerve. 
Calyx prominently 5-winged 
6, Calyx not winged or only very slightly winged (8. amatlana) 


B. beguinotii 
=quidistant or closer to the adjacent lateral nerves 


i 


B. petiolata 


ium 
B. arborea 


B. brachycalyx 


B. geniculata 


WUNDERLIN 


7. Ovary glabrous or glabrat 
8. ee yellow; yeaa in asciulat clusters, often appearing before 
e leaves; Mexico to Costa Ric 
8. oe other than yellow; tower in racemes or panicles, appearing 
well after the leaves; Ecuador o 
9, Leaves 1-4 cm long, piicbed to ca. 3/4 the blade length; fruit a 5 
cm long; Peru 


B. cookii 


weberbaueri 


9. Leaves 5-13. cm ae bilobed to ca. 1/2 the blade length; fruit . 
25 cm long; Ecuado 
10. Corolla white; cial sheath ca. 1/8 the stamen length 
B. seminarioi 
10. Corolla red; staminal sheath 3/5 the stamen length B. haughtii 
7, Ovary tomentose or pilose 
11. Corolla pink, rose, pre with red or pink at the bas 
12. Leaves ovate, 4-10 cm long, 4-7.5 cm wide; carole white with 
red or pink at the base; Colombia and Panama 
12. Leaves suborbicular, 3-4 cm long, 3-4 cm wide; corolla deep 
pink or rose; Ecuador and Peru 
11. Corolla white. 


13. Inflorescence and the lower surface of the leaves brown- 
fe) 


B. picta 


. ayabacensis 


ntose B. seleriana 
13. Inflorescence and the lower surface of the leaves other than 
brown-tomento 
14. Leaf blade : ee 5.5 cm long; petals 2-2.7 cm long; Peru 
B. augusti 


14. Leaf blade 0.8-3 cm long; petals ca. 1 cm long; Mexico. 
15. Leaf blade 2-3 cm long, bilobed to 1/2 the length 


B. andrieuxii 
15. Leaf blade 0.8-1.5 cm long, bilobed nearly to the base 


.amatlana 


Li ae amatlana Wunderlin, sp. nov. (Fig. 1). Type: MEXICO. Oaxaca: ca. L.5 km 

of San Cristobal Amatlan, ca. 80 km S of Oaxaca and 20 km E of (San Andrés) Miahuatlan 

ass 16°19.6'N, 96°22'W), ca. 1,500 m, 8 Jun 2000, J. Santiago Martinez s.n. (HOLOTYPE: USF: 
ISOTYPES: BRIT, F MO, NY). 


A Bauhinia andrieuxii Hemsley foliis minoribus bilobis prope basin differt. 


Shrub; branches brown-tomentose when young, soon glabrate. Leaves with the 
blade chartaceous, 5-7-nerved, broadly suborbicular, broader than long, bilo- 
bate nearly to the base, 0.8-1.5 cm long, 1.2-2.2 cm wide, the innermost pair of 
lateral nerves equidistant or closer to the adjacent lateral nerves than to the 
mid-nerve, the base truncate to slightly cordate, the apex of the lobes rounded, 
the upper surface glabrous, the lower surface sparsely subappressed, short-pu- 
bescent, the petiole 4-6 mm long, brown-tomentose; stipules ovate-lanceolate, 
ca. | mm long, caducous; adpetiolar intrastipular excrescence enlarged, form- 
ing a flattened subulate projection ca. 1 mm long, the others minute. Inflores- 
cence terminal, l- to 3-flowered, the peduncle or rachis strigose; floral bud lin- 
ear-lanceolate, 10-12 mm long, strigose, the apex usually with free calyx lobes 


BRIT.ORG/SIDA 22(1) 


UNIVERSITY OF 
SOUTH FLORIDA 
HERBARIUM 


PLANTS OF MEXICO 


HOLOTYPE 


Marti 


8 


UNIVERSITY OF SOUTH FLORIDA 
HERBARIUM 


) 


Fic. 1. Bauhinia amatlana (holotype, USF 


WUNDERLIN, ACVIOIVIN UE 101 


to 0.5 mm long, sometimes slightly winged and contorted, the pedicel 5-7 mm 
long; bract and bracteoles lanceolate, ca. 0.5 mm long. Hypanthium turbinate, 
1-15 mm long; calyx spathaceous at anthesis; petals white, 10-12 mm long, 
slightly spreading, short-clawed, glabrous, the blade broadly elliptic, 5-6 mm 
wide, the margin entire or slightly crisped, the claw 1-2 mm long; fertile sta- 
mens 10, the outer five 7-8 mm long, the inner ones slightly shorter, the fila- 
ments short-connate at the base, forming a staminal sheath with laciniate pro- 
jections extending 1-2 mm above the sheath, the sheath short-pilose on the inner 
surface, glabrous or glabrate on the outer, the anthers linear-oblong, 1-2 long, 
sparsely pilose; gynoecium 5-7 mm long, the gynophore, ovary, and style 
subequal, brown-tomentose. Fruit not seen. 


Paratype. MEXICO. Oaxaca: ca. 1.5 km NE of San Cristobal Amatlan, ca. 80 km S$ of Oaxaca and 20 
km E of (San Andrés) Miahuatlan (ca. 16°19.6'N, 96°22'W), ca. 1,500 m, 29 Aug 1995, J. Santiago 
Martinez s.n. (USF). 

Etymology.—The name is derived from the Spanish name of the village San 
Cristobal Amatlan, this in turn derived from Amatla, the Aztec name for the 
place of the tree (Ficus sp.) from which the bark is used to make paper. 

Distribution and ecology.—Mexico (Oaxaca). Known only from the type 
locality. Deciduous thorn scrub at ca. 1,500 m. 

Bauhinia amatlana is closely related to B. andrieuxii, also of Oaxaca, but 
differs by its leaf blade 0.8-1.5 cm long and bilobed nearly to the base (vs. 2-3 
cm long and divided only to the middle in B. andrieuxii). 

2. Bauhinia andrieuxii Hemsley, Diagn. Pl. Nov. Mexic. 48.1880; Biol. Cent-Amer, 


Bot. 1:336. 1880. Type: MEXICO. OAXACA: near Oaxaca, - 1834 (FD), G. Andrieux 411 (H¢ 
LOTYPE: K; IsoTYPES: F-fragment, G, K, P; photo ex G: F MO, US). 


Shrub or small tree; branches tomentose when young, soon glabrate. Leaves with 
the blade chartaceous, 7-9-nerved, suborbicular or broader than long, bilobate 
tol/2 the length, 2-3cm long, 2.5-3cm wide, the innermost pair of lateral nerves 
equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the 
base cordate to rounded, the apex of the lobes rounded, the upper surface gla- 
brous, the lower surface tomentellous, the petiole 5-15 mm long, tomentellous; 
stipules ovate, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence 
enlarged, forming a subulate projection ca. | mm long, the others minute. Inflo- 
rescence terminal, racemose, 3- to 8-flowered, the rachis strigose to tomentellous; 
floral bud linear-lanceolate, 12-18 mm long, strigose, the apex with free calyx 
lobes ca. 1.5mm long, the pedicel ca. 5 mm long; bract and bracteoles lanceolate, 
ca. 1.5mm long. Hypanthium cupulate, 2-3 mm long; calyx spathaceous at an- 
thesis; petals white, ca. 1 cm long, slightly spreading, subsessile, glabrous, the 
blade broadly elliptic, 4-6 mm wide, the apex rounded, the margin entire; fer- 
tile stamens 10, the outer five ca. 1 cm long, the inner ones ca. 5 mm long, the 
filaments unequally connate near the base for 1-3 mm, glabrous, the anthers 


102 BRIT.ORG/SIDA 22(1) 


linear-oblong, ca. 2 mm long on the longer stamens, ca. 1.5 mm long on the 
shorter ones, glabrous; gynoecium ca. | cm long, the gynophore, ovary, and style 
subequal, tomentose. Fruit linear, 8-10 cm long, ca. 1.5 cm wide, apiculate with 
the persistent style, dark brown, tomentose, the gynophore 5-10 mm long; seeds 
not seen. 

Distribution and ecology.— Western Mexico (Morelos, Oaxaca, and Sinaloa). 
Deciduous thorn scrub at elevations from near sea level to ca. 1,700 m. 

Bauhinia andrieuxtii is closely related to B. amatlana, also of Oaxaca, but 
differs by its leaf blade 2-3 cm long, bilobed to the middle (vs. 0.8-1.5 cm long, 
bilobed sa to the base in B. amatlana). 


eae 


Specimens examined. MEXICO. Morelos: Canon de Lobos, 14 km on Cuernavaca-Yautepec road, 
Fryxell 764 (MEXU);, Canon de Lobos, Vazquez 1776 (MEXU). Oaxaca: without precise locality, 
Andrieux 371(G, P); Monte Alban, Rose & Hough 4590; Mitla, Smith 107 (US); ruins at Monte Alban, 
Sousa et dl. 6077 (kK, MEXU, MO, WIS); 3 km NE of La Ollaga, Téllez & Magallanes 233 (kK). Sinaloa: 
Cerro Llano Redondo, W of Caimanero, Gentry 7018 (NY). 


3. Bauhinia arborea Wunderlin, sp. nov. (Fig. 2). ee ee BDOR Napo: Anangu, Parque 
Nacional Yasuni, 260-350 m, 30 May-21 Jun 1982 (fD, SE li lorean Forest ]8935 
(HOLOTYPE: AAU; ISOTYPES: AAU, N 


A Bauhinia brachycalyx Ducke foliis majoribus, fructibus latioribus, staminum vagina glabra, nec 
] LaF, 


10N g\ 
Tree to 30 m; branches glabrous. Leaves with the blade coriaceous, (3-)5-7- 
nerved, elliptic-oblong, unlobed, 12-30 cm long, 6-13 cm wide, the innermost 
pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to 
the mid-nerve, the base rounded to obtuse, the apex abruptly short-caudate, 
the upper surface glabrous, the lower surface minutely strigulose, slightly lighter 


in color than the upper surface, the petiole 2-4.5 cm long, glabrous; stipules 
broadly lanceolate, ca. 1 mm long, caducous; adpetiolar intrastipular excres- 
cence enlarged, forming a laterally flattened, obtuse projection to | mm long, 
the others minute. Inflorescence subterminal, short-racemose, 10- to 18-flow- 
ered, the flowers usually paired, the rachis strigose; floral bud elliptic-lanceolate, 
1.5-2 cm long, strigose, the apex acute, the pedicel 1-3 mm long; bract and 
bracteoles triangular-ovate, ca. | mm long. Hypanthium short-tubular, 7-10 mm 
long; calyx spathaceous at anthesis; petals white, 12-15 mm long, slightly 


spreading, sessile, glabrous, the blade oblanceolate, 5-7 mm wide, the apex ob- 
‘ive 12-14 mm 


—— 


tuse, the margin slightly crisped; fertile stamens 10, the outer 
long, the inner ones slightly shorter, the filaments free to the base or very short- 
connate, the anthers lanceolate, 4-6 mm long, sparsely short-pilose on the dor- 
sal surface; gynoecium ca. 8 mm long, the gynophore and style ca. 2 mm long, 
tomentose, the ovary ca. 5 mim long, tomentose. Fruit elliptic-oblong, ca. 13 ¢m 
long, 3-4 cm wide, apiculate with the persistent style base, brown, glabrous, 
C 


— 
— 


ae gynop hore 5-8 cm long; seeds not seen. 


WUNDERLIN, REVISION OF 103 


A 


STUDIES OF ECUADOREAN FORESTS 


; ORR, 
é IX Se Baul rea rhi 

mu ’ 

F * Richard P. Wu 

a University of South Florida = ie , = aan waded Co Duta The New ark 

Se L985 s by Crarina A nad, axi tow Botancal (nathyie, Unwerety of ABrhus 


Fic. 2. Bauhinia arborea (isotype, NY). 


BRIT.ORG/SIDA 22(1) 


Pararyers. ECUADOR. Morona-Santiago: 15 kim Nol Taisha(02°32'S.77°43W), Ortega 96 (USF). Napo: 
Reserva Etnica Huaorani, highway and pipeline “Maxus” under construction (O0°51'S, 76°26'W), 
Aulestia & Andi 938 (MO, USF); Parque Nacional Yasuni, trail toward Rio Daymi (00°52'S, 76°5'W), 
Ceron 3371 (MO, USF); ca. 30 km NNW of Coca, Rio Huashito, Pennington 10592 (kK); Anangu, Parque 
Nacional Y: i 31-32'S, 76°23'W), SEF 8762 (AAV), SEF 8834 (AAU), SEF 8867 (AAU), SEF 8922 
_ago Agrio Canton, Reserva Cuyabeno, Tarapoa-Tipischa, Rio 


(Ay a SEF 10235( AAU). Sucumbios: | 
C ossing (76°15'W, 00°00'S), Palacios etal. 8917 (USF). 


} 


Etymology.—The name is derived from the tree habit remarkable for its size 
within section Amaria in Ecuador. 

Distribution and ecology.—Eastern Ecuador (Napo, Morona-Santiago, and 
Sucum bios). Rain forest on well-drained hilly ground at 250-450 m. 

The species is most closely related to B. brachycalyx but differs by having 
larger leaves, wider fruits, a glabrous staminal sheath, and a tomentose gyno- 
ecium. Bauhinia arborea is a tree to 30 m while B. brachycalyx is a shrub or 
small tree to 6 m, rarely to 15m. 

4. Bauhinia augusti |larms, Repert. Spec. Nov. Regni Veg. 18:234. 1922. Type: PERU, 
RCA: near the confluence of the Rio Chinchipe and Rio Maranon, 700-800 m, | May 


CAJAMAR 
roTyPe: F here designated to replace destroyed B holotype: 


1912 GD, A. Weberbauer 6224 (LEC 

ISOLECTOTYPES: F-fragment, G, NY, US; photo ex B: ER MO, NY, US). 
Shrub to 2m; branches pubescent when young, soon becoming glabrate. Leaves 
with the blade chartaceous, 7-9-nerved, broadly ovate, bilobed to ca. 1/2 the 
length, 3.5-5.5 cm long, 3-5 cm wide, the innermost pair of lateral nerves equi- 
distant or closer to the adjacent lateral nerves than to the mid-nerve, the base 
cordate, the apex of the lobes obtuse, the upper surface glabrous, the lower sur- 
face short-pilose, especially on the nerves, the petiole 0.5-1.5 cm long, short- 
detiolar 


pilose; stipules triangular-lanceolate, ca. 1.5 mm long, caducous; ad 
intrastipular excrescence occasionally enlarged, forming a subulate projection 
to 2 mm long, the others minute. Inflorescence terminal or subterminal and 
axillary, short-racemose or paniculate, 4+- to 12-flowered, the rachis densely 
short-pilose; floral bud narrowly ellipsoid, slightly arcuate, 2-2.5 cm long, short- 
pilose, the apex obtuse or acute, the pedicel 0.5-1 cm long; bract and bracteoles 
triangular-lanceolate, ca. | mm long. Hypanthium short-tubular, +-6 mm long; 
calyx spathaceous at anthesis; petals white, 2-2.7 cm long, slightly spreading, 
short-clawed, glabrous, the blade narrowly elliptic-oblanceolate, 0.5-0.7 cm 
wide, the apex obtuse, the margin slightly crisped, the claw 3-4 mm long; fer- 
tile stamens 10, the outer five 3-3.5cm long, the inner ones slightly shorter, the 
filaments short-connate at the base, sparsely pilose, especially toward the base, 
the anthers elliptic-oblong, ca. 5 mm long, glabrous; gynoecium 3-3.5 cm long, 
the gynophore, ovary, and style subequal, the ovary densely short-pilose, ne 
style and the gynophore sparsely mn Fruit not seen. 
Distribution and ne (Cajamarca). Known only 
collection. At 700-800 m. 


— 


— 


rom the type 


WUNDERLIN, REVISION OF BAUHINIA SUBGENUS BAUHINIA SECTION AMARIA ae 


Bauhinia augusti is related to the new species B. ayabacensis of Ecuador 
and Peru. Bauhinia augusti differs by its white corolla and lower leaf surface 
with erect trichomes in contrast to the pink corolla and lower leaf surface with 
appressed trichomes of B. ayabacensis. 


5. Bauhinia ayabacensis Wunderlin, sp. nov. (Fig. 3). Typr: PERU. Piura: on road to 
Ayabaca, 18 km above Puente Tandopa (Rio Quiroz), 1,700 m, 24 Sep 1964 (fl), PC. Hutchinson 
& J.K. Wright 6689 (HOLOTYPE: F; IsSOTYPES: K, MO, NY, P. US). 
A Bauhinia augusti Harms corolla rosea lata, foliorum lamina inferiori trichomatibus brevibus 
adpressis obsita differt. 
Shrub or small tree to 4 m; branches pubescent when young, soon becoming 
glabrate. Leaves with the blade chartaceous, 7(-9)-nerved, suborbicular, bilobed 
to ca. 1/3 the length, 3-4 cm long, 3-4 cm wide, the innermost pair of lateral 
nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, 
the base cordate, the apex of the lobes obtuse to rounded, the upper surface 
glabrous, the lower surface appressed short-puberulent, especially on the nerves, 
the petiole 1-15 cm long, pubescent; stipules triangular-lanceolate, ca. 1.5 mm 
long, caducous; adpetiolar intrastipular excrescence occasionally enlarged, 
forming a subulate projection to 2 mm long, the others minute. Inflorescence 
terminal or subterminal and axillary, short-racemose, 2- to 10-flowered, the 
rachis brown-tomentose; floral bud narrowly ellipsoid, slightly arcuate, 2-2.5 
cm long, brown-tomentose, the apex acute, the pedicel 6-8 mm long; bract and 
bracteoles triangular-lanceolate, ca.1 mm long. Hypanthium short-tubular, 4- 
6 mm long; calyx spathaceous at anthesis; petals deep pink or rose, 2.5-3 cm 
long, spreading, short-clawed, glabrous, the blade narrowly elliptic-obovate, l- 
L.5cm wide, the apex apiculate, the margins slightly crisped, the claw 3-4 mm 
long; fertile stamens 10, the outer five 2-2.5cm long, the inner ones 1-2 cm long, 
the filaments short-connate at the base, sparsely pilose, the anthers elliptic- 
oblong, ca. 5mm long, glabrous; gynoecium 2-2.5cm long, the gynophore, ovary, 
and style subequal, the gynophore and ovary tomentose, the style sparsely pi- 
lose. Fruit linear-elliptic, ca. 6 cm long, ca. 1.7 cm wide, apiculate with the per- 
sistent style base, brown, strigose to glabrate, the gynophore ca. lcm long; seeds 
ovate-reniform, ca. | cm long, 7-8 mm wide, the surface dark brown, dull, the 
funicular aril-lobe scars unequal, 1-3 mm long. 


Paratypes. ECUADOR. Loja: 5 km E of Celica on road to El Empalme, Harling & Andersson 18183 (GB, 
MO); El Empalme-Celica road, ca. 5 km above El Empalme, Harling & Andersson 18221 (MO). PERU. 
Cajamarca: on the crossing to San Luis, Cabanillas & Guevara 509 (F); El Camote, Carretera Chilete- 
Contumaza Sdagdstegui & Lopez 7636 (F MO, USF); between Rupe and Contumaza Sagdstegui et al. 
9817 (MO); near San Pablo, Sagdstegui et al. 15380 (F); Caserio de Chilili, Sanchez 200 (F); between 
Chilete and Rupe, E of the Chilete-Contumaza highway, Sdnchez 4222 (F). 


Etymology.—The name derived from the town of Ayabaca (Province Ayabaca) 
near the type locality. 


BRIT.ORG/SIDA 22(1) 


UNIVERSITY OF CALIFORNIA 


Director, Paul C. Hutchison 
Additional field sores for fag 
and 3m. in diame 


: Tre 
r. Flowers deap pink. 


Irwin, 1966, comments ae description 
\ gives color as white." 
Bauhinia sages std Harm 
dupl. de STiadin; wes 
HOLOTYP 
Sie ayabacensis Wunderlin Dry open hillsides, scattered brash On road to Ayabaca, 
R4199r 18 km. : 
1641236 rch. Wen ie en 
niversity of South Florida 1982 Biovioce of iArabaes: ficschae Hi ae 
FIELD MUSEUM OF Pau C. Hutcuison 7, 
NATURAL HISTORY Collection number 6689 of veh po on 1, UCy J. Kenneth Wr T 24 September 1964 
determiners set, NY s Seaees USM, US, F, NY, MO, K, C 
MICH, M, P, LE, 


Fic. 3. Bauhinia ayabacensis (holotype, F) 


WUNDERLIN, REVISIUN UF 107 


Distribution and ecology.—Ecuador (Loja) and Peru (Cajamarca and Piura). 
Dry, open hillsides in scattered brush or seasonal evergreen forest at 1,200-2,400 
m. 

Bauhinia ayabacensis is closely related to B. augusti from which it differs 
by its wider deep pink- or rose-colored corolla (vs. white in B. augusti) and the 
lower leaf surface with appressed trichomes (vs. short pilose in B. augusti). The 
petals of B. ayabacensis are 1-15 cm wide, while those of B. dugusti are 0.5-0.7 
cm wide. 

6. Bauhinia beguinotii Cufod., Arch. Bot. Sist. 9:192. 1933. Type: COSTA RICA. LIMON: 
between Waldeck and Rio Barbilla, 40 m, 12 May 1930 (fD, G. Cufodontis 664 (HOLOTYPE: W, 
not seen; photo ex W; ISOTYPES: F MO, US). 


Shrub or small tree to 6(-10) m; branches brown-tomentose to -tomentellous 
when young, soon becoming glabrate. Leaves with the blade chartaceous, 7-9- 
nerved, oblong-ovate, unlobed or slightly bilobate (deeply bilobate or bifoliolate 
in var. gorgonae), 10-24 cm long, 5-16 cm wide, the innermost pair of lateral 
nerves closer to the mid-nerve than to the adjacent lateral nerves, the base 
rounded to truncate, the apex Gif the leaf unlobed) or the lobes (if the leaf bi- 
lobed or bifoliolate) acuminate to caudate, the upper surface glabrous, the lower 
surface brown-tomentellous to glabrate, the petiole 1.5-3.5 cm long, glabrate; 
stipules broadly ovate, 1-1.5 mm long, caducous; adpetiolar intrastipular ex- 
crescence occasionally enlarged, forming a subulate projection to 1 mm long, 
the apex glandular, the others minute. Inflorescence terminal or subterminal 
and axillary, racemose, 5-10-flowered, the rachis brown-tomentose to - 
tomentellous; floral bud elliptic-lanceolate, 1.5-2 cm long, brown-tomentose, 
the apex with minute free calyx lobes, the pedicel 2-5 mm long; bract and 
bracteoles ovate, ca. 1 mm long. Hypanthium short-tubular, 5-6 mm long; ca- 
lyx spathaceous at anthesis; petals white, 2.5-3 cm long, spreading, short-clawed, 
glabrous, the blade oblanceolate, 8-13 mm wide, rounded, the margin entire, 
the claw 3-5 mm long; fertile stamens 10, the outer five 2-3 cm long, the inner 
ones shorter, the filaments connate for ca. 1/2 the length of the shorter stamens, 
sparsely pilose near the base, the anthers triangular at anthesis, ca. 3 mm long 
on the longer stamens, ca. 2 mm long on the shorter ones, sparsely pilose to 
glabrate; gynoecium 2-3 cm long, the gynophore, ovary, and style subequal, 
brown-tomentose. Fruit linear, 12-15 cm long, ca. 2 cm wide, apiculate with the 
persistent style, brown, minutely strigose to glabrate, the gynophore 1-1.5 cm 
long; mature seeds not seen. 


KEY TO THE VARIETIES OF BAUHINIA BEGUINOTII 


1. Leaves unlobed or only slightly bilobate; Costa Rica and Panama var. beguinotii 
1. Leaves deeply bilobate or bifoliolate; Colombia var. gorgonae 


108 BRIT.ORG/SIDA 22(1) 


6a. Bauhinia beguinotii var. beguinotii 


Distribution and ecology.—Costa Rica (Limon) and Panama (Bocas del Toro, 
Panama, and San Blas). Tropical evergreen forest from near sea level to ca. 500 m. 
Specimens examined. COSTA RICA. Limon: drainage of the Rio Estrilla Valley, Shank & Molina 4466 
(fF, US). PANAMA. Bocas del Toro: Cerro Bracha, NE of the encampment Changuinola | of Corriente 
Grande (IRHE), Correa et al. 3227 (PMA, USF); SE and NE of the encampment Changuinola | of the 
IREIE, Correa et al. 3316 (PMA, USF). Panama: 14.8 kin N of the Pan-American Highway, Folsom & 
Maas 5202 (MO); Carti road, 7 mi from turnoff at El Llano (9°15'N, 78°50'W), Hammel 13547 (MO. 
USF): 10-12 km N of Pan-American Highway, Mori & Kallunki 2847 (MO), El Llano-Carti-Tupile 
road, 16 km N of Pan-American Highway at El Llano, Nee 9362 (MO): ca. 10 km N of Pan-American 
Highway, Nee etal. 8881(MO). San Blas: FE] Llano-Carti road (09°20'N, 79°O'W), Hamilton & Stockwell 


COCK WELL 


1089 (MO, USF): El Llano-Carti road, 19.1 km (09°19'N, 78°55'W ). de Nevers 5946 (MO, USF): ena 


along continental nes on E L L ke Carti sauce a“ LO'N, 78°15'W), de Nevers & Pérez 3701 ( 
JSF) ). Nusagandi | g the E aes Carti road (O9°19'N, 78°15'W), de — 
& Pérez 3582; Nusagandi g tl tal d El Llano-Carti road (09°1L9'N, 78°15'W), 


de Nevers & Pérez 3781 (MO, USP): Nusangandi ridge running NW from Punta Mama, (09°19'N, 
78°15S'W), de Nevers et al. 3728 (MO, USF); El Llano-Carti road, 19.1 km from Interamerican | 
(O9°19'N, 78°55'W), de Nevers et al. 4957 (MO, USF); El Llano-Carti road, near Nusagandi, aor 
Sendero Nusagandi, W of buildings (09°15'N, 79°O'W), McPherson 11033 (MO, USF). 


6b. Bauhinia beguinotii var. gorgonae (Killip ex R.S. Cowan) Wunderlin, Ann. 
Missouri Bot. Gard. 60:570. 1973. Bauhinia gorgonde Killip ex R.S. Cowan, Bol. Soc. 
Venez. Ci. Nat. 22:281. 196L. Type: COLOMBIA. NARINO: E side of Isla Gorgona, 50-100 m, 11 
Feb 1939 ({D, EP. Killip & H. Garcia-Barriga 33170 (HOLOTYPE: US; ISOTYPES: COL, not seen, 
BM, US). 

Distribution and ecology.—Colombia (Narino). Tropical evergreen forest from 

near sea level up to 100 mm. 

The Colombian material is recognized at the varietal level and is 
distinquished from the Panamanian and Costa Rican material only in having 
the leaves deeply bilobate or bifoliolate rather than unlobed or only slightly 
bilobate. Since the degree of lobing of leaves in Bauhinia is often quite variable 
within species, further collections may prove var. gorgonde to be unworthy of 
recognition. 


Specimens examined. COLOMBIA. Narino: Isla Gorgona, Cheeseman [St. George Expedition] 380 (F K, 
US), Collenette [St. George Expedition] 628 (EK, US), Kelsall [St. George Expedition] 360 (K, US). 


7. Bauhinia brachycalyx Ducke, Trop. Woods 90:12. 1947. Typr: BRAZIL. AMAZONAS: 
region of the Rio Purus near the mouth of the Rio Pauini [based on plant cultivated in the 
Horto Botanico do Museu Goeldi, Belém, Para, specimen made 30 Dec 1946 (I1)|, A. Ducke 
2045 (HOLOTYPE: MG, not seen; IsOTYPES: NY, US). 

Shrub or small tree to 6(-15) m; branches glabrous. Leaves with the blade sub- 

coriaceous, 5-nerved, oblong-ovate, unlobed, 7-11 cm long, 3-5.5 cm wide, the 

innermost pair of lateral nerves equidistant or closer to the adjacent lateral 

nerves than to the mid-nerve, the base rounded to subtruncate, the apex witha 

caudate tip 1-2 cm long, the upper surface glabrous, the lower surface glabrous, 


— 


WUNDERLIN, REVISION Ur 109 


the petiole 0.8-1.4 cm long, glabrous; stipules ovate, ca. | mm long, caducous; 
adpetiolar intrastipular excrescence often enlarged, forming a subulate projec- 
tion to 2 mm long, the others minute. Inflorescence terminal or subterminal 
and axillary, short-racemose, 6- to 12-flowered, the flowers usually paired, the 
rachis minutely puberulent; floral bud elliptic-oblanceolate, ca. 1.5 cm long, 
minutely puberulent, the apex apiculate, the pedicel 3-4 mm long; bract and 
bracteoles ovate-lanceolate, ca. | mm long. Hypanthium short-tubular, 4-6 mm 
long; calyx spathaceous at anthesis; petals white, ca. 1.5(-2) cm long, slightly 
spreading, subsessile, glabrous, the blade linear-oblanceolate, 4-5 mm wide, the 
apex acute, the margins slightly crisped; fertile stamens 10, the outer five 8-10 
mm long, the inner ones ca. 1/2 as long, the filaments short-connate at the base, 
forming a staminal sheath with a laciniate projection extending 1-2 mm above 
the sheath, the sheath tomentose on the inner surface, glabrous on the outer, the 
anthers linear-oblong, ca. 3 mm long, sparsely pilose at the apex and the base; 
gynoecium 12-15 mm long, subequaling the androecium, arcuate, the ovary 5-6 
mm long, the gynophore and the style 3-4 mm long, subequal, the style and the 
upper portion of the ovary with glandular trichomes in lines, the gynophore gla- 
brous, the stigma oblique. Fruit linear, 10-12 cm long, 1.5-2.5 cm wide, apiculate 
with the persistent style, brown, glabrous, the gynophore ca. 0.5 mm long; seeds 
suborbicular, 11-12 mm long, 9-10 mm wide, dark brown, the surface dull, the 
funicular aril-lobe scars subequal, extending ca. 1/2 way around the seed. 

Distribution and ecology.—Ecuador (Napo and Sucumbios), Peru 
(Amazonas, Loreto, and Pasco), and Brazil (Amazonas). In forest on well drained 
ie also often on inundated clay soil and along rivers at 200 m. 


pecimens examined. ECUADOR. Morona-Santiago: oil well “Garza” of Tenneco, ca. 35 km NE of 
Mere Ivo (01°49'S, 76°42'W), Zak & yee als 4618 (MO, USF) Napo: road construction on the Yuca 
Road, ca. 3 km from the Auca oil-field Road (ca. 00°31'S, 76°58'W,) Brandbyge et al. 30287 (AAU, 
USF); Parque Nacional Yasuni, oil well “bogi” of Conoco (00°43'S, 76°28W), Coello 225 (MO, USF); 
Anangu, N side of Rio Napo, trail from village to laguna (00°31'S, 76°23'W), Lawesson et al. 39571 
(AAU, USF); Anangu, near the outlet of Rio Anangu into Be a in pus vues Nacional Yasuni, 
SEF 10133 (AAU); Anangu, near the outlet of Rio At | Yasuni, 


SEF 10389 (AAU). Sucumbios: Rio Gueppi, tributary ee Rio preys above Beauv border post of 
Puerto Pert, 8 km above mouth of river, Gentry et al. 21805 (MO, USF); Lago Agrio Canton Reserva 
Faunistica Cuyabeno, Rio Lagarto, on border with Peru (00°33'S, 75°16'W), Palacios et al. 8055 (MO, 
USF). PERU. Amazonas: Yamayakat, Quebrada Kusu-Chapi, Rio Maranon (00°55'S, 78°19W), Vasquez 
et al. 20104 (MO, USF). Loreto: Chambira, near Yurimaguas, Rio Huallaga Valley, Ferreyra 4952 (NY, 
US); Rio Gueppi, tributary of Rio Putumayo, northernmost tip of Peru on border with Ecuador, be- 
tween mouth of River and the border a of Puerto Pertica. 8 km upriver, Gentry et al. 21772 (MO, 
USF); trail from Indi iana on Rio Amazonas to Rio Napo, Gentry et al. 22188 (MO, USF); Rio Nanay on 
the road tothe M f Be ne rista, Rimachi 2800 (MO); San Miguel (Quebrada Tamshiyacu) 
ae a 72°40'W), Vasquez & Jaramillo 7153 (MO, USF); unos) saeaee a Suerte (Rio Itaya), Vasquez & 

8418(MO, USF): Indiana, Yanamono, Rio A S,72°50'W), Vasquez & Jaramillo 
‘11103 (MO, USP): Indiana, Yanamono, Explorama Lodge (03°28'S, 72°50 W), Vasquez et al. 12113 (MO 
USF). Pasco: Palcazu Valley, Iscozacin (10°12'S, 75°15'W), Foster 9463 (MO, USF). BRAZIL. Amazonas: 
Rio Javari, Miraflor, Lleras et al. [Prance] P16927 (K, MO, US) 


110 BRIT.ORG/SIDA 22 


1) 


8. Bauhinia cookii Rose in Britton @ Rose, N. Amer. FL. 23:205. 1930. Type: COSTA 
RICA, ALAJUFLA: Cuesta de la Vieja, road to San Carlos, 300 m, 11 Apr 1903 (1D, O.F Cook & 
CB. Doyle 41 (HOLOTYPE: US; IsoTYPES: NY-fragment, US 


Casparia calderonii Rose in Britton & Rose, N. Amer. Fl. 23:217.1930. Ba ‘alderonii (Ros se) 
Lundell, pollen 1214. 1937. Type: EL SALVADOR, SONSONATE: San Julian, 1924 (Ir), S 
Calderén 2226 (HOLOTYPE: US; ISOTYPES: NY-fragment, US). 

Bauhinia gigas L oT Lyra 1:213. 1937. TYPE: ee Cayo: Valentin, Jun-Jul 1936 (st), 
CL. Lundell MICH; IsoTyPEs: NY, US). 


Tree to 15(-40) m; branches strigose or puberulent when young, soon becom- 
ing glabrate. Leaves with the blade submembranaceous or chartaceous, 7-9- 
nerved, broadly ovate, bilobate to ca. 1/2 the length, 5-10(-15) cm long, 7-10(- 
12) cm wide, the innermost pair of lateral nerves equidistant or closer to the 
adjacent lateral nerves than to the mid-nerve, the base cordate to truncate, the 
apex of the lobes acute to obtuse, the upper surface sent the lower surface 
strigose, at least on the nerves, the petiole 2-3(-5) cm long, slightly strigose; 
stipules lanceolate, ca. 1.5 mm long, caducous; adpetiolar intrastipular excres- 
cence enlarged, forming a subulate projection to 1.5 mm long, the others minute. 
Inflorescence paniculate, congested into a fasciculate cluster, sometimes pro- 
duced before the leaves and appearing cauliflorous or ramiflorous, 15- to 60- 
flowered, the rachis strigose; floral bud linear-elliptic, 8-10 mm long, strigose, 
the apex with minute free calyx lobes, the pedicel 5-7 mm long, strigose; bract 
and bracteoles ovate-lanceolate, ca. | mm long. Hypanthium cupulate, 2-3 mm 
long; calyx spathaceous at anthesis; petals light sulfur-yellow, 10-15 mm long, 
spreading, sessile or short-clawed, glabrate, the blade elliptic to oblanceolate, 
5-7 mm wide, the apex obtuse, the margin entire; fertile stamens 10, the outer 
five LO-15 mm long, the inner ones 5-10 mm long, the filaments connate at the 
base 
linear-oblong, ca. 2 mm long on the longer stamens, ca. 1.5 mm long on the 


or ca. L mm, with a laciniate rim projecting above, glabrate, the anthers 


shorter ones; gynoecium 5-10 mm long, the gynophore, ovary, and style 
subequal. Fruit linear, 5-7 cm long, ca. 1.5 cm wide, apiculate with the persis- 
tent style, light brown, See the gynophore 5-10 mm long; seeds obovate, 
7-10mm ee 5-8 mm wide, the surface dull, dark brown, the funicular branch 
scars equal, ca. 4/5 the length of the seed. 

Distribution and ecology.—Western Mexico in Sinaloa southeast through 
Mexico and Central America to Costa Rica. Not known from Nicaragua, but to 


be expected there. Tropical deciduous or semideciduous forests at elevations 
from near sea level to ca. 4,200 m, but more commonly 300-1,000 m. It appar- 
ently flowers throughout the year, but usually from March to July. Flowers are 
sometimes produced before the leaves appear or are much developed. 

The species is unusual not only in section Amaria by having yellow flow- 


— 
—_ 


ers, but in fact, no other New World species of subgenus Bauhinia has flowers 


of this color. 


WUNDERLIN, REVISION Ur 111 


Specimens examined. MEXICO. Chiapas: vicinity of La Chacona, Miranda 6485 (US). Colima: vicin- 
ity of Manzanillo, Ferris 6022 (A, F US); ca. 15 mi SSW of Colima on Manzanillo road (3 mi above 
Terolapa), Mc Vaugh 15541 (MICH); ca. 14 mi WNW of Santiago on road to Cihuatlan, oe isco, Mc a 
20776 (G, MICH, US). Guerrero: Canon de la Mano Negra, near Iguala, Rose et al. 9344 (F GH, NY, US). 
Jali ween Bahia Navidad and La Manzanilla on Bahia Tenacatita, 3 mi W of ee hae 
road, McVaugh a 007 (MICH); near Playa de Cuastecomate, 8 km by road NW of Navidad, McVaugh 
& Koelz 1674 (MICH). Nayarit: 7-8 mi W of Compostela along road to Mazatlan, Gentry & Gilly 
10848 (DUKE, MEXU, TEX, WIS); 6-7 mi S of Compostela, McVaugh 18753 (G, MICH, US). Oaxaca: 
Ejido Benito Juarez, Sebastopol, Tuxtepec, Dioscorea Brigade 2741 (MEXU). Sinaloa: Mazatlan, Reko 
4520 (MEXU, US). Veracruz: Barranca de ae Purpus 8797 (UC), Purpus 8896 (GH, MO, NY, UC, 
Uo) near Rancho Remundadero, Purpus 11068 (NY, US), Purpus 11069 (A), Purpus 11170 (MO); 
Municipio Puente Nacional, La Ceiba, Ventura 12760 (MEXU). GUATEMALA. Escuintla: Hacienda 
ae Salas 1406 (F); Rio Guncalate Standley 60187 (F, US). Guatemala: without precise locality, 
Aquilar 610 (F). Retathuleu: Rio Talculan, 5 mi W of Retalhuleu, Standley 87377 (F). Suchitepéquez: 
along Rio Madre Vieja Joa oe ae 62207 (F). HONDURAS. Comayagua: Pito Solo, Lake 
Yojoa, Edwards P-404 (Fk, US). EL DOR. Ahuachapan: Finca San Benito, near Rio Guayapa 
(13°49'N, 89°56'W), aoe et al. oon U sea Finca San Bene alone Rio Nacimiento del Guayapa 
(13°5 EN, 89°55'W), Witsberger Nts COSTA RICA. Al etween Canas and Upala, 
10 km N of Pyaepe Croat 36478 (MO, US nee of the Rio Sue aes Pittier 16695 (US). San José: 
Cerro Bijaqualito-Tarcoles, Poveda 1097 es 


9. Bauhinia geniculata Wunderlin, sp. nov. (Fig. 4). Type COLOMBIA. RISARALDA: Haci- 
enda Alejandria, 6 km on highway between La Virginia and Cerrito, low hills of the extreme 
north of the wide part of the Valle del Rio Cauca, ca. 940 m, 27 May 1989 (f1), PA. Silverstone- 

Sopkin, J. Giraldo, & M. Cabrera 5270 (HOLOTYPE: CUVC; ISOTYPE: USF) 


A Bauhinia petiolata (DC.) Triana ex Hooker f. foliis lobatis, calyce alto, nec non androecii forma 
singulari differt. 

Tree to 5 m; branches glabrous. Leaves with the blade coriaceous, 7-9-nerved, 
oblong-ovate, bilobed to ca. 2/3 the length, 14-26 cm long, 9.5-17 cm wide, the 
innermost pair of lateral nerves equidistant or closer to the adjacent lateral 
nerves than to the mid-nerve, the base shallowly cordate to subtruncate, the 
apex of the lobes acuminate, the upper surface inconspicuously reticulate, gla- 
brous, the lower surface evidently reticulate, with inconspicuous, colorless, 
medially attached, biradiate, glandular trichomes, minutely strigose, at least 
on the veins near the base, the petiole 2-3(-4) cm long, glabrous; stipules trian- 
gular-ovate to lanceolate, 1-1.5 mm long, caducous; adpetiolar intrastipular ex- 
crescence sometimes enlarged and forming a laterally flattened, blunt projec- 
tion to 2 mm long, others minute. Inflorescence terminal or subterminal and 


axillary, racemose, 2- to 10-flowered, the flowers solitary or paired, the rachis 
reddish brown-tomentose; floral bud linear-clavate, 4-4.5 cm long, sparsely 
brown-tomentose, evidently 10-nerved, becoming prominently 5-winged above 
the hypanthium with the wings gradually enlarging toward the apex to 4mm, 
the pedicel ca. 1.5cm long; bract and bracteoles triangular-ovate, ca. 1 mm long. 
Hypanthium short-tubular, L.5-2.2 cm long; calyx green, spathaceous at anthe- 
sis, each sepal with a thickened nectiferous area ca. 5-7 mm long and wide near 
the base; petals white, 3.5-4.5 cm long, slightly spreading, subsessile, glabrous, 


112 BRIT.ORG/SIDA 22(1) 


UNIVERSITY OF 
SOUTH FLORIDA 
HERBARIUM 


k 


Fic. 4. Bauhinia geniculata (isotype, USF). 


WUNDERLIN, ACVIOIUIN UE 113 


the blade oblanceolate, 15-18 mm wide, gradually tapering to the thickened 
midrib at the base, the apex rounded to slightly g : fertile stamens 10, 
the outer five 4-4.5 cm long, the inner ones shorter, the filaments connate at 
the base for 2-3 mm and forming a staminal sheath, the uppermost stamen of 
the inner whorl the shortest and strongly geniculate near the point of attach- 
ment near the apex of the shield-like portion of the staminal sheath, the stami- 
nal sheath with a ligulate projection ca. 1mm long behind the lower 3 stamens 
of the outer whorl and with a minute projection behind the lower 2 stamens of 
the inner whorl, the sheath reddish tomentose on the inner surface, the anthers 
linear, 7-8 mm long, glabrous; gynoecium 3.5-4.5 cm, the gynophore ca. 1.5 cm 
long, reddish tomentose, the ovary ca. 2 cm long, reddish tomentose, the style 
ca. 15cm long, reddish tomentellous to glabrate near the apex. Fruit linear, 29- 
41 cm long, 2.8-3.3 cm wide, apiculate with the persistent style base, brown, 
puberulent, the gynophore 2.5-4 cm long; seeds suborbicular to oblong, strongly 
compressed, 17-18 mm long, 12-15 mm wide, the surface dull brown, the fu- 
nicular scars subequal, extending 2/3 to nearly the length of the seed. 


ParatyPe. COLOMBIA. Risaralda: Hacienda Alejandria, 6 km on highway between La Virginia and 
Cerrito, low hills of the extreme north of the wide part of the Valle del Rio Cauca, Silverstone-Sopkin 
et al. 5809 (CUVC, USF). 
Etymology—The name derived from the bent uppermost stamen of the inner 
staminal whor 
Distribution and ecology—Colombia (Risaralda). Primary forest at 900-940 m. 
Bauhinia geniculata is most closely related to B. petiolata, but differs by its 
lobed leaves (vs. unlobed in B. petiolata), winged calyx (vs. not winged in B. 
petiolata), and unique androecial morphology. 


10. Bauhinia haughtii Wunderlin, Brittonia 35:337. 1983. Type: ECUADOR. MANaBi: 
Bella Vista, SE of Santa Ana, 100 m, 5 Sep 1942 (f1, fr), O. Haught 348] (HOLOTYPE: US; ISOTYPES: 


Shrub or tree to 10 m; branches glabrous. Leaves with the blade submembrana- 
ceous, 9(-11)-nerved, suborbicular, bilobed to ca. 1/2 the length, 5-8 cm long, 6- 
10 cm wide, the innermost pair of lateral nerves equidistant or closer to the 
adjacent lateral nerves than to the mid-nerve, the base deeply cordate, the apex 
of the lobes obtuse to rounded, the upper surface aac the lower surface 
minutely strigose, the petiole 3-5 cm long, glal iangular-lanceolate, 
ca. l mm long, caducous; intrastipular ¢ excrescences ‘minute. Inflorescence ter- 
minal or subterminal and axillary, racemose, 6- to 10-flowered, the flowers usually 
paired, the rachis minutely strigulose-tomentose; floral bud narrowly elliptic, 
5-6 cm long, minutely strigulose-tomentose, red, the apex attenuate, the pedicel 
2-5 mm long: bract and bracteoles triangular-lanceolate, ca. 1 mm long. Hy- 
panthium tubular, 2-2.5cm long, slightly arcuate; calyx spathaceous at anthe- 
sis; petals red, 4.5-5cm long, slightly spreading, short-clawed, glabrous, the blade 


114 BRIT.ORG/SIDA 22(1) 


oblanceolate, 1-1.5 cm wide, the apex rounded, the margin entire, the claw 3-4 
mim long; fertile stamens LO, red, the outer five 4.5-5.5 cm long, the inner ones 
slightly shorter, 9 connate into a staminal tube for ca. 3/5 their length, the other 
one free to ca. 0.5 cm lower and slightly longer, the staminal sheath sparsely 
short pilose on the inner surface, the anthers elliptic-oblong, ca. 5 mm long, 
glabrous; gynoecium 6.5-8.5 cm long, the gynophore, ovary, and style subequal, 
glabrous. Fruit linear, 16 cm long, 1.7 em wide, apiculate with the persistent 
style, light brown, glabrous, the gynophore 4.5 cm long: seeds not seen. 

Distribution and ecology.—Ecuador (Manabi and Pichincha). In tropical wet 
forest at L1OO-300 m. 

Bauhinia haughtii is a distinctive species most closely related to B. 
seminarioi, also endemic to Manabi, Ecuador. It differs from that species by its 
red flowers (vs. white in B. seminarioi) and a much longer staminal sheath (ca. 
3.5.cm vs. ca.6 mm in B. seminarioi). The only other red-flowered species in 
Ecuador, B. stenantha, is readily distinguished by its five fertile stamens (vs. 
ten) and smaller flowers (2.5-3.0 cm long) and smaller fruits (6-10 cm long). 


Specimen examined. ECUADOR. Manabi: 170-175 km on road between Santa Domingo and Quininde, 
Acosta-Solis 13653 (F). 


11. Bauhinia petiolata (DC.) Hook. f., Bot. Mag. 103: tab. 6277. 1877. Amaria petiolata 
DC., Prodr, 2: 519. 1825. Type: COLOMBIA: without precise locality, 1760-1808, 5. Mutis 2398 


(HOLOTYPE: MA, not seen; ISOTYPE: US). 

Amaria sessilifolia DC, Prodr. 2:519, 1825. Bauhinia sessilifolia (DC.) Quinones, Ann. Missouri 
Bot. Gard. 75:1156. 1988. TyPE: COLOMBIA: without precise locality, 1760-1808, S. Mutis 2724 
(HOLOTYPE: MA, not seen; Isotype: US-fragment). 


ane 


Shrub or tree to 15 m; branches glabrous. Leaves with the blade chartaceous to 
coriaceous, 5-nerved, ovate, unlobed, 8-14 cm long, 4-8 cm wide, the innermost 
pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to 
the mid-nerve, the base rounded to deeply cordate, the apex acute, sometimes 
with a caudate tip to 3. cm long, the upper surface glabrous, the lower surface 
glabrous, often glaucous, conspicuously reticulate-nerved, the petiole 0.5-3.5 
cm long, glabrous; stipules triangular, ca. 1 mm long, caducous; adpetiolar 
intrastipular excrescence often enlarged and forming a blunt subulate projec- 
tion ca. | mm long, the others minute. Inflorescence terminal or subterminal 
and axillary, short-racemose, 4- to 8-flowered, the rachis sparsely strigose- 
tomentellous to glabrate; floral bud elliptic-ovate, 2-3 cm long, strigulose-to- 
mentose, the apex attenuate, glabrous, the pedicel to 1 cm long; bract and 
bracteoles triangular-ovate, ca. 1 mm long. Hypanthium short-tubular, ca. 1.5 
cm long; calyx spathaceous at anthesis; petals white, 3.5-4.5 cm long, slightly 
spreading, short-clawed, glabrous, the blade narrowly oblanceolate, 1.2-1.5 cm 
wide, the apex rounded, the margin slightly crisped, the claw 2-3 mm long: 
tertile stamens 10, the outer five 3-5(-7.5) cm long, the inner ones slightly shorter, 


—" 


WUNDERLIN, AEVISIUN UF 115 


the filaments short-connate at the base, glabrous, the anthers linear-oblong, 8- 
12 mm long, glabrous; gynoecium 4-5 cm long, the gynophore ca. 1 cm long, 
glabrous, the ovary ca. 3 cm long, tomentose (glabrous or glabrate in var. 
caudigera), the style ca. 1.5cm long, glabrous. Fruit linear, 21-27 cm long, 2-2.5 
cm wide, apiculate with the persistent style, brown, sparsely pubescent, gla- 
brescent at maturity, the gynophore ca. 2.5 cm long; seeds not seen. 

Bauhinia petiolata is a highly distinctive though poorly known species 
represented by two varieties. The species is most closely related to B. geniculata 
which is also from the upper Magdalena Valley (Risaralda Prov.) in Colombia 
but is readily distinguished by its unlobed leaves and flower buds not winged. 
Italso has affinities with B. arborea (Ecuador) and B. brachycalyx (Ecuador, Peru, 
and Brazil), two other species with unlobed leaves in section Amaria. 

Quinones (1988) argues that B. sessilifolia is a distinct species and differs 
from B. petiolata by its sessile leaves, the leaf base deeply cordate, the inflores- 
cence of axillary umbels, the petals emarginate, the stamens of five long and 
five short and included, and the fruit 2 cm wide. Bauhinia petiolata on the other 
hand has leaves with the petiole 0.5-3 cm long, the leaf base truncate or rounded, 
the inflorescence a terminal or axillary in a short raceme, the petals acute, the 
stamens subequal and excluded, and the fruit 1.5 cm wide. These characters, 
with the exception of the sessile leaves, do not hold up in the material exam- 
ined. Even though I have not seen the MA material cited by Quinones, I believe 
that B. sessilifolia is based on aberrant, sessile-leaved specimens of B. petiolata. 
The sessile-leaved condition is approached in some material seen for B. petiolata. 
For example, André 1810 has petioles only 5 mm long. 


KEY TO THE VARIETIES OF BAUHINIA PETIOLATA 


1. Ovary tomentose; central Colombia var. petiolata 
1. Ovary glabrous or glabrate; Panama and Venezuela var. caudigera 


lla. Bauhinia petiolata var. petiolata 
Distribution and ecology.—Upper Magdalena Valley of central Colombia 
(Cundinamarca and Tolima). Wet forest at ca. 500 m. 


Specimens examined. COLOMBIA. Cundinamarca: Quebrada Cachimbulo near Tocaima, Rio Funza 
valley, André 1810 (kK); Rio Magdalena valley, Goudot s.n. (P); Hacienda El Cucharo, between Tocaima 
and Pubenza, Killip et al. 38314 (COL, FE NY, US), Killip et al. 38350 (COL, K, US); between Nilo an 
Puebla Nuevo, Murillo & Jaramillo 260 (COL, NY); Tocaima, Triana 4412 (BM, COL, G, NY); La Parada 
Limba and the Rio Magdalena valley, Triana s.n. (P); Tocaima, Triana s.n.(K); Tocaima, Rio Magdalena, 
Triana s.n. (kX). Tolima: Chicoral, Haught 6306 (COL, NY, US). State unknown: without locality, Mutis 
2377 (fragment ex MA: US); Mutis 2549 (fragment ex MA: US); Mutis 4146 (fragment ex MA: US). 
Cultivated at Kew from material provided by Linden from Colombia (K). 


11b. ue petiolata var. caudigera (S.F Blake) vi peeiete comb. nov. BASIONYM: 
a caudigera S.F. Blake, Contr. U.S. Natl. Herb. 20:522. 1924. TYPE: VENEZUELA. 
Baie ales, alone road from Puerto Cabello to os Felipe, 10-100 m, 20 May-10 


BRIT.ORG/SIDA 22(1) 


Jun, 1920 ({D, H. Pittier 8851 (HOLOTYPE: US; photo ex US: US, NY, ISOTYPE: P). 
Distribution and ecology.—Panama (Colén) and Venezuela (Carabobo and 
Falcon). Rainforest and wet forest at 10-600 m. The variety was probably more 
widely distributed from Panama through northern Colombia to Venezuela in 
the past. Its present disjunct distribution is relictual, apparently now confined 
to wet river valley refugia. 

This taxon was placed in synonymy of B. petiolata by Wunderlin (1983). 
Since then, additional material has become available and it is here considered 
as a distinct variety on the basis of its glabrous or glabrate ovary (vs. tomentose 
in var. petiolata). 


} 


Specimens examined. PANAMA. Colon: ca. 2-3 mi up the Ri mouth, Kennedy & Fos- 
ter 2127 (MO, USF). VENEZUELA. Carabobo: upper Ciaremales ve from Puerto Cabello to San Fe- 
lipe, Pittier 8914 (NY, US). Faleon: Parque Nacional Quebrada de la Cueva E1 Toro (L0°50'N, 69°07'W), 


Liesner etal. 7727 (MO. USF), Liesner et al. 7736 (MO), Wingfield 7501 (USF). 


12. Bauhinia picta (Kunth) DC,, Prodr. 2: O15. 1825. ale etia picta RUE, in a Eisibolie! 
al., Nov. Gen. Sp. 6:316. 1824. Type: COLOMBIA. ! A g the Ri near the 
confluence with Rio Opon, between Bojorque and Isla de etree May 1801 (fD), RWH.A. 
Humboldt & AJ.A. Bonpland 1604 (HOLOTYPE: P, not seen, microfiche IDC 156/C6; ISOTYPEs: P, 

5). 


BCW) not seen; photo ex BCW): F G, MO, NY 


Bauhinia ligulata Pittier, Contr. U.S. Natl. Herb. 20:112. 1918. Type: PANAMA. SAN BLAS: near Pu- 
erto Obaldia, 0-50 m, Aug 1911 ({L), H. Pittier 4334 (HOLOTYPE: US; IsoTYPES: BM, F NY, US). 
Seana kalbreyeri Harms, Repert. Spec. Nov. Regni Veg. 19:65. 1923. TYPE: COLOMBIA. 

ANTIOQUIA: Murri, 1,000-1,100 m, Jun 1880 (£D, W. ees 1802 (LECTOTYPE: K, here ae 

nated to replace paves B holotype; ISOLECTOTYPES: F-fragment, US; photo ex K: EF NY 

photo ex B: EK, S 
Tree to 20(-40) m; branches ferruginous-tomentose when young, soon becom- 
ing glabrate. Leaves with the blade chartaceous, 9-13-nerved, ovate, bilobate to 
ca. 1/4 the length, 4-10(-21) cm long, 4-7.5(-15) cm wide, the innermost pair of 
lateral nerves equidistant or closer to the adjacent lateral nerves than to the 
mid-nerve, the base cordate to rounded, the apex of the lobes apiculate to obtuse 
or rounded, the upper surface glabrous, the lower surface finely ferruginous- 
tomentellous (at least on the nerves), the petiole 2-3.5 cm long, ferruginous- 
tomentellous to glabrate; stipules ovate, ca. 1 mm long, caducous; adpetiolar 
intrastipular excrescence enlarged, forming a subulate projection to 2 mm long, 
the apex with a lateral glandular area, the others minute. Inflorescence axil- 
lary, subterminal, short-racemose, 10- to 20-flowered, the flowers solitary or 
paired, the rachis ferruginous-tomentose; floral bud linear-clavate, 2-2.5 cm 
long, ferruginous-tomentose, the apex with minute free calyx lobes, the pedicel 
2-6 mm long; bract and bracteoles lanceolate, ca. 1mm long. Hypanthium short- 
tubular, 6-8 mm long; calyx spathaceous at anthesis, soon splitting partly or 
fully to the hypanthium into 2 to 5 lobes; petals white with pink or roseate 
base, sometimes lavender-pink or rose, 2-3 cm long, spreading, short-clawed, 


WUNDERLIN, ACVIOIVUN UE 117 


glabrous, the blade ovate-elliptic, 6-13(-20) mm wide, the apex rounded, the 
margin entire, the claw 2-4 mm long; fertile stamens 10, the outer five 2-3 cm 
long, the inner ones slightly shorter, the filaments free to base or short-connate, 
short-hirsute or with only a few short trichomes on the lower 1/2, the upper 
stamen attached near the apex or the middle of the ligulate part of the staminal 
sheath, short-hirsute (rarely glabrate); gynoecium 2-3 cm long, the gynophore, 
ovary, and style subequal, the gynophore sparsely tomentose below the ovary 
on the lower surface or glabrate, the ovary tomentose, the style sparsely 
tomentellous to glabrate. Fruit linear, 15-25 cm long, ca. 2 cm wide, apiculate 
with the persistent style, brown, tomentellous to glabrate, the gynophore 1-2 
cm long; seeds elliptic-ovate, ca. 15 mm long, ca. 12 mm wide, dark brown, the 
surface dull, the funicular aril-lobes 6-7 mm long. 

Distribution and ecology.—Colombia (Antioquia, Bolivar, Boyaca, Choco, 
Cundinamarca, Risaralda, and Santander) and Panama (San Blas). In open to 
closed forest from near sea level to about 1,500 m. 

The species is apparently most closely related to B. petiolata, but differs in 
having bilobed leaves and petals white with a pink or roseate base. The type of 
B. ligulata from Panama isa large tree (40 m) with slightly smaller flowers and 
the adaxial stamen attached lower on the ligulate portion of the staminate 
sheath. It agrees in all other respects with the Colombia material. The material 
from Choco Province, Colombia differs by its slightly larger leaves (16-21 cm 
long compared with the usual 4-10 cm long), the glabrate staminal sheath and 
filament bases (these usally short-hirsute), and the less tomentose ovary. 

Bauhinia picta is occasionally cultivated in Colombia for its showy flowers. 


1. COLOMBIA. Antioquia: 14-17 km N of Remedios on the way to Zaragoza, Cerro 

Cakes aa along the Rio Tucupé (07°20N, 74°30'W), Callejas et al. 5177 (USF); Vereda Alicante, 
Finca Penjamo, Quebrada Penjamo, on the San Juan de Bedout-La Cabana road (06°39'N, 74°31 W), 
Callejaset a 9388 (HUA nee Vereda La Pava, 14 km by Andes highway toward La Vereda El Libano, 
Fonnegra et al. 2441 (USF); Segovia, Sandeman 5575 (K). eee : near the confluence of 
the Rio Ité and Rio Tamar into the Rio Cimitarra, ca. 38 km W of mn eee ja (06°55 'N, 74°15'W), 
- Br a 1554 (MO, NY, US). Boyaca: fl Umbo region, ie rence 492 (K). ae Rio Tagachi, ca. 12 
of Rio Atrato (O6°LSN, 76°50'W), Gentry et al. 37068 (MO, USF); Mecana, Juncosa 1923 (MO, 


= we Cundinamarca: 16 km NW of Alban along highway to Villeta, Barclay et al. 3666 (US). Risaralda: 
Hacienda ee ndria, 6 km on La Virginia-Cerritos highway, extreme northern part of Valle del Rio 
Cauca, Silverstone-Sopkin 5895 (CUVC, USF). Santander: Puerto Berrio, between Rio Carare and Rio 
Magdalena, ee 1772 (P); Bucaramanga and vicinity, Killip & ain 14978 (NY). State unknown: 
Rio Carare, Dawe 456 (K); Cordillera Orientale, Goudot s.n. (P). 


13. Bauhinia seleriana Harms in Loes., Bull. Herb. Boissier 7:549. 1899. Type: GUA- 
AL A rae JETENANGO: near Quen Santo, ca. 1,100 m, C. Seler & E. Seler 2797 (LECTOTYPE: 
K,h stroyed B holotype; ISOLECTOTYPE: F-fragment, NY, US; photo 
ex K: E NY, US: cae EG, MO, NY US:). 
Bauhinia paradisi Standley & L.O. Williams, Ceiba 1:80. 1950. TyPpE: HONDURAS. EL PARAISO: 
Rio Lizapa floresta de Robles area, between Galeras and Lizapa Grande, 1,000 m, 14 May 1948 
(fl, A. Molina 860 (HOLOTYPE: US; photo ex US: NY, US; ISOTYPES: BM, F). 


118 BRIT.ORG/SIDA 22(1) 


Shrub or small tree to 12 m; branches short-hirsute when young, soon becom- 
ing glabrate. Leaves with the blade chartaceous, 9-11-nerved, broadly ovate, bi- 
lobate to ca. 1/3 the length, 6-10 cm long, 6-10 cm wide, the innermost pair of 
lateral nerves equidistant or closer to the adjacent lateral nerves than to the 
mid-nerve, the base cordate to rounded, the apex of the lobes obtuse to rounded, 
the upper surface glabrous, the lower surface short hirsute or tomentellous, the 
petiole (I-)2-4(-5) cm long, hirsute or tomentellous; stipules triangular-ovate, 
ca. 4mm long, caducous; adpetiolar intrastipular excrescence often enlarged 
and forming a subulate projection to 1.5 mm long, the others minute. Inflores- 
cence terminal or subterminal and axillary, racemose, 5- to 30-flowered, the 
rachis short-hirsute or tomentellous; floral bud elliptic, 8-15 mm long, short- 
hirsute or tomentellous, the apex with free lobes to 2 mm long, the pedicel 4-5 
mm long; bract and bracteoles lanceolate to linear, 1-4 mm long. Hypanthium 
cupulate, 1-2 mm long; calyx spathaceous at anthesis; petals white, 11-18 mm 
ong, spreading, subsessile or short-clawed, glabrous except for the blade apex, 
the blade elliptic, 4-8 mm wide, the apex apiculate, the margin entire, the claw 
ca. 1 mm long, glabrous; fertile stamens 10, the outer five 11-18 mm long, the 
inner ones 1/2 the length of the outer, the filaments unequally connate at the 
base for 1-3 mm, with a laciniate rim extending above the point of adnation, 
the inner surface tomentose, the anthers linear-oblong, ca. 2 mm long on the 
longer stamens, ca. 1.5 mm long on the shorter; gynoecium 11-18 mm long, the 
gynophore, ovary, and style subequal, tomentose. Fruit linear, 8-10 cm long, ca. 
15cm wide, apiculate with the persistent style, dark brown, tomentose, the gy- 
nophore 0.5-1 cm long; seeds not seen. 

Distribution and ecology.—Southern Mexico, Guatemala, and Honduras. 
Deciduous forest at 700-1,500 m. 

The species is closely related to B.andrieuxii of western Mexico from which 
itis Gennes by its larger leaves 6-10 cm long (vs. 2-3 cm in B. andrieuxii) 
and conspicuously hirsute parts. Bauhinia seleriana is a forest species while B. 
andrieuxti occurs in open deciduous scrub. 


—_— 


Representative specimens examined: MEXICO. Chiapas: along Mexican Highway 190 3 mi S of La 
Trinitaria, Breedlove 14485 (NY). México: gan ancho, Hinton etal. a One ca. paces 


EF of Laollaga, Téllez & Magallanes 233 (MEXU, MO). GUATEMALA. I 
a7 Las Palmas, ne eee 51582 (F US). ane Loma El Picacho, above Santa Rosalia Steyermark 
42730 (F NY, US). H RAS. Choluteca: 6 km SW of Panamerican Highway, Harmon & Fuentes 


5962 (UMO). El Paraiso: ie Rio California, Sierra de la Villa Santa, Williams & Molina 10479 (F, 
MO, UC, US, VT). Francisco Morazan: along Quebrada Suyapa, near Suyapa, Molina 553 (F& MO, US). 


14. Bauhinia seminarioi Harms ex Eggers, Bot. Centralbl. 73:69. 1898. Type: ECUA- 
DOR. MANABI near mecienda El pec 16 mae ee (fl, fr), HEA. von Eggers 14925 (LECTO- 
YPE: F-fragment, h d B holotype; photo ex B: ISOLECTOTYPES: 

FE a US). 


Shrub or small tree 2-3(-5) m; branches sparsely tomentellous when young, 


WUNDERLIN, REVISION OF 119 


soon becoming glabrate. Leaves with the blade submembranaceous to 
chartaceous, (5-)7-9-nerved, suborbicular, bilobed to ca. 1/2 the length, 5-13 
cm long, 6-12 cm wide, the innermost pair of lateral nerves equidistant or closer 
to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex 
of the lobes obtuse to rounded, the. upper surface glabrous, the lower surface 
minutely strigulose, short-pilose along the veins, especially near the base, the 
petiole 2-4.5 cm long, tomentellous, soon becoming glabrate; stipules triangu- 
lar-lanceolate, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence 
occasionally enlarged and forming a subulate projection to 2mm long, the oth- 
ers minute. Inflorescence terminal or subterminal and axillary, racemose, 8- to 
18-flowered, the flowers usually paired, the rachis tomentellous to glabrate; flo- 
ral bud linear-clavate, 4-6 cm long, tomentellous to glabrate, the apex rounded, 
the pedicel 0.5-1.8 cm long; bract and bracteoles triangular-lanceolate, 0.5 mm 
long. Hypanthium short-tubular, 1.5-2 cm long; calyx spathaceous at anthesis, 
petals white, ca. 5cm long, slightly spreading, short-clawed, glabrous, the blade 
linear-oblanceolate, 5-6 mm wide, the apex acute to obtuse, the margin slightly 
crisped, the claw ca. 0.5cm long; fertile stamens 10, the outer five ca. 4 cm long, 
the inner ones slightly shorter, the filaments short-connate at the base to ca. 6 
mim, the staminal sheath pilose on the inner surface near the apex and slightly 
above on the filaments, the anthers elliptic-oblong, ca. 5 mm long, glabrous; 
gynoecium 5.5-6.5cm long, the gynophore, ovary, and style subequal, glabrous. 
Fruit linear, 15-25 cm long, 2-2.3 cm wide, apiculate with the persistent style, 
dehiscent, brown, glabrous, the gynophore 2-3 cm long; seeds not seen. 

Distribution and ecology.—Ecuador (Guayas and Manabi). Tropical moist 
forest on inner coastal hills at 100-150 m. 

The type collection (von Eggers 14925) is apparently a mixture of this spe- 
cies and Bauhinia aculeata subsp. grandiflora Jussieu ex Poiret) Wunderlin of 
subg. Bauhinia section Bauhinia. Specimens of B. aculeata subsp. grandiflora 
with various collection dates were distributed as the type of B. seminarioi by C 
to K, F and undoubtedly elsewhere. The description, except for the statement 
that the species is aculeatous, clearly fits what is known to be B. seminarioi as 
represented by the photograph of the holotype (now destroyed) and not B. 
aculeata subsp. grandiflora. The adpetiolar intrastipular excrescences in B. 
seminarioi sometimes enlarge to 2 mm, especially in the inflorescence, but the 
older branches can not be described as aculeatous. Since the holotype is of young 
floriferous material with evident enlarged excrescences, Harms apparently in- 
correctly interpreted the materia. The Iltis & Iltis E-221 specimen bears only 
small blunt excrescences. 


mens ined. ECUADOR. eal Bosque Protector Cerro Blanco, road to Salinas, 15 km 
(02° 10'S, 79°58'W), Rubio 2020 (MO, USF). Manabi: Hacienda El Recreo, von Eggers 14886 (F, K, P, US); 
4 km E of Hacienda El Recreo (2 as S along coast from Canoa, 18 km due N of Bahia de Caraquez; 
00°27'S, 80°26'W) and the Pacific Ocean, Iltis & Iltis E-221 (USF WIS) 


120 BRIT.ORG/SIDA 22(1) 


15. Bauhinia webert i Harms, Bot. Jahrb. Syst. 42:91. 1908. Type: PERU. CAJAMARCA: 
near eas 2 sia 2,100 m, Jan 1905 (fD, A. Weberbauer 4802 (LECTOTYPE: G, here desig- 
nated t stroyed B holotype; ISOLECTOTYPE: F-fragment; photo ex B: F MO). 


Bauhinia ruiziana Harms, Repert. Spec. at Regni Veg. 18:234. 1922. Type: PERU. HUANUCO: 

erie. io ee ay H. Rune aoe 1795 [H. Ruiz Lopez & J.A. Pav6én 1300] (LECTOTYPE: K, 
erate ISOLECTOTYPES: F, MA, not seen, P; photo ex K, 
F US; Tes MA F nets ex B. Fi G MO, U 


Shrub to 1.5m; branches strigulose when young, soon becoming glabrous. Leaves 
with the blade chartaceous, 7-nerved, suborbicular, bilobed to ca. 3/4 the length, 
15-4 cm long, 15-4 cm wide, the innermost pair of lateral nerves equidistant 
or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, 
the apex of the lobes obtuse to rounded, the upper surface glabrous, the lower 
surface strigulose, the petiole 0.5-1 cm long, minutely strigulose or glabrous; 
stipules triangular-lanceolate, |-1.5 mm long, caducous; adpetiolar intrastipular 
excrescence occasionally enlarged and forming a subulate pone to2mm 
long, the others minute. Inflorescence terminal or subterminal, the flowers soli- 
tary, the rachis strigulose; floral bud elliptic-lanceolate, ca. 2 cm long, red, 
strigulose, the pedicel ca. 1 cm long; bracts and bracteoles triangular, ca. | mm 
long. Hypanthium short-tubular, 4-6 mm long; calyx spathaceous at anthesis: 


petals bright pink or rose-red, 3-4 cm long, spreading, short-clawed, glabrous, 
the blade oblanceolate, 1-1.8 cm wide, the apex rounded, the margin entire, the 
claw 3-5 mm long; fertile stamens 10, red, the outer five 2.5-3.cm long, the inner 
ones slightly shorter, the filaments very short-connate, sparsely pilose toward 
the base, the anthers elliptic-oblong, 3-4 mm long, glabrous; gynoecium 3-4 
cm long, the gynophore, ovary, and style subequal, glabrous. Fruit narrowly el- 
liptic, 4.5-5 cm long, ca. 1.5 em wide, apiculate with the persistent syle, dehis- 
cent, light brown, glabrous, the gynophore 4.5-5 cm long; seeds hee tees y el- 
liptic-ovate, dark brown, the surface dull, the funicular aril-lobe scars subequal, 
2-3 mm long. 

Distribution and ecology.—Peru (Cajamarca, La Libertad,and Huanuco). On 
steep rocky slopes above rivers at 2,100-2,400 m. 

Bauhinia weberbaueri is related to B. ayabacensis and B. dugusti, but is eas- 
ily distinguished from both by its glabrous ovary (vs. tomentose in B. augusti 
and B. ayabacensis) and from B. augusti by its pink to rose petals (vs. white in B. 
d soci 


fo 


PERU. La Libertad: between Los Alisos and Pataz, Alayo 18 (USF). Huanuco: 
above eiasenda bute, between Ambo and Huanuco, Ferreyra 6553 (F); Ambo, Macbride & 
Featherstone 2411 (F), Macbride 3164 (FG), Pavon s.n. (BM), Sawada P-103 (F); Huanuco, Machride 
3503 (FG), Matthews 927 (kK), without precise locality, Vita, McLean s.n. (K). 


Il. Series Stenanthae Wunderlin, K.Larsen & S.S.Larsen. Biol. Skr. 28: 14. 1987. 


Type: Bauhinia stenantha Diels 


Petals strongly imbricate, the corolla forming a pseudotube; fertile stamens 5. 


WUNDERLIN, REVISION OF 121 


1. Bauhinia stenantha Diels, Biblioth. Bot. 29(116):93. 1937. Type: ECUADOR, 
CHIMBORAZO: canyon of the Boo panciae near Paes each ] ie m, 7-14 May 1945 (£1, fr), 
W.H. Camp E-3041 (NEOTYPE: Fh B holotype; ISONEOTYPE: 
K, P). [ECUADOR. CHIMBORAZO: en of the Rio c shanchan near Huigra, 1,260 m, 21 Sep 
1933 (fl, fr), EL.E. Diels 1130 (HOLOTYPE: B, destroyed)]. 


Shrub or small tree to 3 m; branches puberulent when young, soon becoming 
glabrate. Leaves with the blade submembranaceous or chartaceous, 7-9-nerved, 
suborbicular, bilobed to ca. 3/4 the length, 2.5-5.5 cm long, 3-5.5 cm wide, the 
innermost pair of lateral nerves equidistant or closer to the adjacent lateral 
nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse to 
rounded, the upper surface glabrous, subglaucous, ns LOWE sue sparsely to 
moderately tomentose, the petiole 1-3cm long g te; stipules 
triangular-ovate, ca. 1mm long, caducous; adpetiolar intrastipular excrescence 
occasionally enlarged and forming a subulate projection to 2 mm long, the oth- 
ers minute. Inflorescence terminal or subterminal, short-racemose, 2- to 1O- 
flowered, the rachis tomentellous or strigulose; floral bud elliptic-oblanceolate, 
ca. 2.5 cm long, rose-red, tomentellous or strigulose, the apex apiculate, the 
pedicel 1-1.5cm long; bract and bracteoles triangular-lanceolate, ca. 1 mm long. 
Hypanthium short-tubular, subgibbous at the base, 6-8 mm long; calyx 
spathaceous at anthesis, soon splitting to the hypanthium into 2 to 5 lobes; pet- 
als rose-red, 2.5-3 cm long, erect and pseudotubular, short-clawed, glabrous, the 
blade oblanceolate, 7-8 mm wide, the apex rounded, the margin slightly crisped, 
the claw 8-9 mm long; fertile stamens 5, red, 2.5-3 mm long, the filaments very 
short-connate, the staminal sheath with a small laciniate rim, glabrous, the 
anthers oblong, ca. 5mm long, glabrous, the staminodes 5, rudimentary or to 1/ 
2 the length of the fertile stamens, the aborted anthers globose; gynoecium 3.5- 
45cm long, the gynophore, ovary, the style subequal, glabrous. Fruit linear, 6- 
10 cm long, ca. 1.5 cm wide, apiculate with the persistent style base, dehiscent, 
light brown, glabrous, the gynophore ca. 1.5 cm long; mature seeds not seen. 

Distribution and ecology.—Ecuador (Azuay and Chimborazo). Dry scrub 
hillsides at (600-)1,200-1,800 m. 

|. ECUADOR. Azuay: along Pasaje-Santa Isabel-Giron highway, ee of the Rio 

Tenens Harling & Andersson 14404 (GB, USF); Cuenca-Machala road, between Girén and Santa Isa- 
bel, 17 km from Girdén (03°14'S, 79°13'W), Lojtnant & Molau 14176 (AAU); 17 om SW of Giron, 
MacBryde 451 (MO). Chimborazo: Huigra, Asplund 7736 (S), Asplund 15507 (K, S); vicinity of Huigra 
Rose & Rose 22119 (photo ex NY: US); Huigra, White s.n. (BM). State unknown: without definite local- 
ity, Acosta-Solis 11675 (F). 


ACKNOWLEDGMENTS 
| gratefully acknowledge the curators of A, AAU, BM, COL, CUVC, DUKE, EG, 
GB, GH, HUA, K, MEXU, MICH, MO, NY, P PMA, S, TEX, UC, UMO, US, VT, WIS 
for making specimens available for study. I thank Philip A. Silverstone-Sopkin, 
(CUVC) for information on Bauhinia geniculata, Gerald G. Robinson (USF), for 


122 BRIT.ORG/SIDA 22(1) 


information on Bauhinia amatlana, Bruce F. Hansen (USF) for his helpful sug- 
gestions on the manuscript, and Kathleen Hotchkiss (USF) for assistance with 
graphics. | also thank the three reviewers of the manuscript for their helpful 
suggestions. 


REFERENCES 


ForTUNATO, R.H. 1986. Revision del genero Bauhinia (Cercideae, Caesalpinioideae, Fabaceae) 
para la Argentina. Darwiniana 27:527-557. 

Fortunato, RH. 1997. Bauhinia. Flora Fanerogamica Argentina. 33:4—9. Programa Proflora 
(CONICET), Cordoba, Argentina. 

McVaucu, R. 1987. Leguminosae. In:W.R. Anderson, ed. Flora Novo-Galiciana. 5:1—786. Uni- 
versity of Michigan Press, Ann Arbor. 

Quinones, L.M. 1988. Una nueva combinacion en Bauhinia (Fabaceae—Caesalpinicideae). 
Ann. Missouri Bot. Gard. 75:1155-1156. 

Vaz, A.M.S.DAF.1979.Consideragdes sobre a taxonomia do género Bauhinia L.sect. Tyloteae 
Vogel (Leguminosae-Caesalpinioideae) do Brasil. Rodriguésia 31: 

Vaz, A.M.S. ba F. and A.M.G. pe A. Tozz!. 2003. Bauhinia ser. Cansenia (Leguminosae: 
Caesalpinioideae) no Brasil. Rodriguésia 54:55—143. 

Wuwoeruin, R.P. 1976. The Panamanian species of Bauhinia (Leguminosae). Ann. Missouri 
Bot. Gard. 63:346-354. 

WUNDERLIN, R.P. 1983. Revision of the arborescent Bauhinias (Fabaceae: Caesalpinioideae: 
Cercideae) native to Middle America. Ann. Missouri Bot Gard. 70:95-127. 

WuNbeatin, R.P. 1998. Bauhinia. In:J.A. Steyermark, PE. Berry, and B.K. Holst, eds. Flora of Ven- 
ezuelan Guayana. 4:5-13. Missouri Botanical Garden Press, St. Louis. 

Wunberun, R.P. 2001. Bauhinia. In: W.D. Stevens, C.U. Ulloa, A. Poole, and O.M. Montiel, eds. 
Flora de Nicaragua. 1:522-526. Missouri Botanical Garden Press, St. Louis. 

WUNDERLIN, R.P., K. LARSEN, and S.S. Larsen. 1987, Reorganization of the Cercideae (Fabaceae: 
Caesalpinioideae). Biol. Skr. 28:1—40. 


NO 
™ 


DENDROPHORBIUM RESTINGAE (ASTERACEAE: 
SENECIONEAE), A NEW SPECIES FROM SAO PAULO, BRAZIL 


Aristonio M. Teles Jimi N. Nakajima 
Universidade Federal de Minas Gerais Universidade Federal de Uberlandia 
Instituto de Bichel ias iB Qeaiea? Rua Ceara s/n - Bloco 2D - campus Umuarama 


Departam Uberlandia, Minas Gerais 
Av. Anténio ete 6627, Pampulha, 31.270-110 
Belo Horizonte, Minas Gerais, BRASIL 
aristonio@hotmail.com 


Joao R.Stehmann 


Universidade Federal de Minas Gerais 
ie tuto ae as éncias Ries 


Ay Antonio ane 6627 
Pampulha, 31.270-110 
Belo Horizonte, Minas Gerais, BRASIL 


ABS RACT 
Dendrophorbium restingae (Asteraceae: Senecioneae), a new species from Sao Paulo state, Brazil, is 
described, illustrated, and compared with the related species D. paranense (Malme) Matzenb. & 
Baptista 

RESUMEN 


Dendrophorbium restingae eeu ee Seceipncae) una nueva epens ae Bae Ee, Brasil, es 
2. | 


descrita, ilustrada y i i aptista 


L 


Dendrophorbium (Cuatrec.) C. Jeffrey is a predominantly Andean genus with 
about 50 species ranging from the Antilles to southern Brazil (Jeffrey 1992). Hind 
(1993) recorded nine species for Brazil; some years later Matzenbacher and 
Baptista (1997) proposed one more new combination in the genus, for a total of 
ten Brazilian species. 

Dendrophorbium was segregated from the large genus Senecio 
(Nordenstam 1996), and is recognized by its large and generally dentate leaves, 
corymbiform to paniculiform capitulescences, radiate heads with 8-13 glabrous 
phyllaries, style branches with a truncate, obtuse, or conical apex bearing a 
tuft of trichomes, and 5-8-ribbed cypselae with an annuliform carpopodium 
(Nordenstam 1996). 


Dendrophorbium restingae A. Teles J. N. .Nakaj. & Stehmann, 2 nov. (Figs. la, b, 
TYPE: BRAZII P gica Juréia-Itatins, restinga 
da praia do Arpoador, 22 Jun 1994, I. Cae M.M.R. Fiuza de Melo, RJ. Oliveira & M. Barros 

1531 (HOLOTYPE: SP; ISOTYPE: K). 


SIDA 22(1): 123 - 128. 2006 


BRIT.ORG/SIDA 22(1) 


124 


lotype, SP). 


a 


' right (HUIULY Pe, 


TELES ET AL., A NEW SPECIES OF DENDROPHORBIUM FROM BRAZIL 125 


INSTITUTO DE BOTANICA sp 
SSo Paulo — Brasil 


Fic. 1b. Basal leaves (holotype, SP). 


BRIT.ORG/SIDA 22(1) 


10 mm 


Fic.2.Dend bi j A. Head. B. Phyllaries.C. Ray floret. D. Disk floret. E. Cypselae of disk floret. F. Stamen. 


All from the holoiyne (SP). 


A enone horbium paranense affinis, sed foliis majoribus vald ‘datis ( 
nulatisq ecules aan pevols non auriculatis, floribus dian 30- 35 (vs beac 25) et 
cypsel is 5-8-costatis (vs. 10 Ca a 


Robust herbs 1-1.5 m high. Stems fistulose, multisulcate and lanate. Leaves al- 
ternate, coriaceous, discolorous, glabrous and brownish adaxially and greyish- 
tomentose abaxially when dry, penninervate with 9-16 secondary veins promi- 
nent abaxially, margins crenulate; basal leaves non-auriculate, petiolate, the 
petioles 5.5-12 cm long, winged, clasping the stem, lanate, the blades ovate-cor- 
date, 7-35 cm long, 6.5-26 cm wide, with apex acute, base cordate; distal leaves 
sessile and invaginating, the blades oblong, 2-14 cm long, 0.7-3.5 cm wide, with 
acute apex and obtuse base. Capitulescences corymbiform. Heads heterogamous, 


TELES ET AL., A NEW SPECIES OF DENDROPHORBIUM FROM BRAZIL 127 


radiate, peduncle 1-4 cm long, lanate, bracteole lanceolate. Involucres |-1.3 cm 
long, 1-1.2 cm wide, campanulate, calyculate, calyculus 5-bracteolate, bracteole 
5-6 mm long, linear, glabrous; phyllaries ca. 13, uniseriate, lanceolate, 10-11 mm 
long, 1.5-2 mm wide, the apex acute and ciliate, the margins glabrous and scari- 
ous. Receptacles plane, alveolate, epaleaceous. Ray florets 4-8, pistillate; corolla 
liguliform to 13 mm long, bidentate to tridentate apically; style to 10 mm long, 
branches straight, ca. 1 mm long, the apex with a tuft of pennicilate trichomes. 
Cypselae 3-4 mm long, cylindrical, 5-ribbed, glabrate, with inconspicuous 
carpopodium; pappus ca. 7 mm long, deciduous. Disk florets perfect, 30-35; 
corolla tubulose, 7-9 mm long, 5-lobate, the lobes 2.5-3 mm long, the midvein 
visible along the distal half of the tube; stamens with anthers to 3 mm long, 
exserted, caudate at base to 0.2 mm long, the filaments with anther collar di- 
lated, the connective with an oblong conspicuous appendage to | mm long; style 
7-12 mm long, branches 1-2.5 mm long, divergent, apex truncate crowned by 
simple trichomes. Oypeclac 3-5 mm long, cylindrical, 5-8ribbed, glabrate, 
carpopodium annuliform; pappus 6-7 mm long, deciduous. 

Dendrophorbium restingae mor phological) resembles Dend rophorbium 
paranense (Malme) Matzenb. & Baptista, but is disti ble by its crenulate 
and larger leaves and more ray florets, as well as its distinct geographical range 
and habitat. The new species so far has been collected only in coastal forests on 
sandy soils of the state of Sao Paulo. This area belongs to the edaphic vegeta- 
tional complex called Restinga that occupies a narrow belt along the Brazilian 
coast (Veloso et al. 1991; Rizzini 1997). Other endemic species of Asteraceae grow- 
ing in this environment, such as species of Noticastrum from Santa Catarina 
and Rio Grande do Sul (Zardini 1985) and Litothamnus from Bahia (Holmes 
1996). Additionally, D. paranense occurs in humid soils of riverine habitats in 
the cloud forests of Parana, Santa Catarina, and Rio Grande do Sul. Both species 
can be distinguished by the characters set in the following key: 


Leaves coriaceous with crenulate margins; petioles not auriculate; phyllaries apically 
ciliate; disk florets 30-35, corollas 7-9 mm long; cypselae of disk florets 5-8-ribbed 
D. restingae 


eee pee earn ql Fee | remot 1 ni- 


alee dik florets 22-25, corollas 3.5- 5n mm long; ees of disk florets . ae 
D. paranense 


Etymology.—The epithet alludes to the habitat of the species, the Brazilian 
coastal vegetation on sandy soils named restinga. Until the present, no species 
of this genus had been collected in this vegetation. 

Geographical distribution and habitat.—This species so far has been col- 
lected only in restinga vegetation at the Juréia-Itatins Ecological Station, in the 
municipality of Peruibe in the state of Sao Paulo, where it grows in rocky places 
along the shore line. 

Phenology.—Flowers and fruits were collected from June to October. 


128 BRIT.ORG/SIDA 22(1) 


PARATYPE: BRAZIL. Sao Paulo: Estacao Ecoldgica de Juréia-Itatins, Municipio de Peruibe, nucleo, praia 
do Arpoador, vegetacao de restinga arbustiva, 18 Oct 2004, Moreno et al. 100 (SP). 


ACKNOWLEDGMENTS 


We thank Inés Cordeiro, curator of the herbarium SP and collector of the type 
material, Marcos Sobral for suggestions and comments on the manuscript, John 
Pruski and Guy Nesom for reviewing the manuscript, Rosa Alves for the line 
drawings, CAPES and CNPq for the grants to the first and last author respec- 
tively. 


REFERENCES 


Hino, DJ.N. 1993. A checklist of the Brazilian Senecioneae (Compositae). Kew Bull. 48: 
279-295. 

Hoimes, W.C. 1996. Litothamnus nitidus (Compositae: Eupatorieae), a new combination 
based on Mikania nitida (DC.) R. King & H. Robinson. Phytologia 81:385-390. 

Jerrrey, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an 
annotated world check-list of the genera of the tribe. Notes on Compositae: VI. Kew 
Bull. 47:49-109. 

Martzensacuer, N.I. and L.R.M. Baptista. 1997. Uma nova combinagdao no género 
Dendrophorbium (Cuatrec.) C.Jeffrey (Asteraceae - Senecioneae). Bol. Mus. Bot. Prefeitura 
Municipal Curitiba, MBM. 65(65):1-3. 

Norbenstam, B. 1996. Recent revision of Senecione and Calenduleae systematics. In DJ.N. 
Hind & HJ. Beentje, eds. Compositae: Systematics. Proceedings of the International 
Compositae Conference. Vol. 1. Kew: Royal Botanic Gardens. Pp. 591-596. 

Rizzini, C.T. 1997. Tratado de fitogeografia do Brasil: aspectos ecoldgicos, socioldgicos e 
floristicos. Ambito Cultural Edicdes Ltda, Rio de Janeiro. 

Vetoso, H.P, A.L.R. Ranaet FitHo, and J.C.A. Lima. 1991. Classificagao da vegetagao brasileira, 
adaptada a um sistema universal. IBGE, Rio de Janeiro. 

ZARDINI, EM. 1985. Revision del genero Noticastrum (Compositae — Astereae). Revista Mus. 
La Plata, Secc. Bot. 13(83):313-424. 


NOVEDADES EN GENTIANACEAE PARA 


AMERICA DEL SUR 
Eva M. Filippa Gloria E. Barboza 
Instituto Multidisciplinario de Instituto Multidisciplinario de 
Biol ogia Vegetal (CONI CET- UNC) y Biologia | vegeldl Vaan T- UNC) y 
Facultad de Ciencias Quimicas (UNC) Facultad de Ciencias fees (UNC) 
Casilla de Correo 495, 5000, Cordoba, Casilla de Correo 495, 5000, Cérdoba, 
ARGENTINA. efilippa@imbiv.unc.edu.ar ARGENTINA. gbarboza@imbiv.unc.edu.ar 
ABSTRACT 


Gentianella fabrisii (Gentianaceae) is proposed as a new species. It is known only from the type 
collection from the northwestern of Argentina (Prov. Tucuman); a key to distinguish this species 
from. its closer eae is provided: ne syuonyas for Cicendia quadrangularis, Curtia tenuifolia, 
4 spe cies o een are proposed. The ocurrence 
of Gietenanthas Vernier 1 Schultesia gui is confirmed. These 
re ] t . Pal : 


fo) oO 


cor 
d and illustrated in detail. 


RESUMEN 


Se describe e ilustra una nueva especie de Gentianella a fa crs) que see en el noroeste de la 


Argentina (Prov. Tucu man), y se provee UNa Clave | 

Se dan a conocer nuevos sinénimos para Cicendia adams Girea tenuifolia, Gentiana 
prostrata, Zygostigma australe y para 4 especies de G nella. Se confirma la pre a de 
Chelonanthus viridiflorus y Schultesia guianensis para : ae y aoe las que se pe 
e ilustran detalladamente. 


INTRODUCCION 


Li tesdl el ti t tartico, 


Gentianaceae es una familia 
con ca. 1690 especies que habitan, por lo comun, en regiones templadas, 
subtropicales y tropicales (Struwe et al. 2002). Una moderna clasificacion en 
tribus y subtribus basada, en gran parte, en estudios filogenéticos derivados de 
datos moleculares (trnL, matK e ITS), acepta 6 tribus y 87 géneros (Struwe et al. 
2002). Esta familia se caracteriza por presentar algunas entidades bien 
conocidas por sus principios amarogénicos y colorantes, de uso tanto en la in- 
dustria alimenticia como en la farmacéutica Jensen & Schripsema 2002). 

En el Nuevo Mundo, la mayor diversidad de géneros (de 47 nativos, 36 son 
endémicos) se encuentra en la region tropical de América Central y América 
del Sur (Struwe et al. 2002). Con motivo de estar realizando la revision de 
Gentianaceae para el Catalogo del Cono Sur y para la Flora Fanerogamica de 
Argentina, surgieron algunas novedades, que ya se dieron a conocer en parte 
(Filippa & Barboza 2001; 2003), a las que se suman ahora las de la presente 
contribucion. 


SIDA 22(1): 129 — 143. 2006 


130 BRIT.ORG/SIDA 22(1) 


UNA NUEVA ESPECIE DE GENTIANELLA 


Gentianella fabrisii Filippa et Barboza, sp. nov. (Fig. 1). Tivo: ARGENTINA. TUCUMAN, 
Dpto. Tafi: Quebrada del Baron, Los Planchones, 3300 m, 7 Feb 1958, H.A. Fabris 1417 a 


(HOLOTIPO: BM; ISOTIPO: H.A. Fabris 1417 a pp., LP). 


A Gentianella bromifolia foliis ovatis vel late ovatis, floribus hermaphroditus et femineis, calycis 


lobis anguste ovatis, corollae lobis tubo 3-5 plo longioribus, g ssili differt. 


Hierba anual, ginomonoica, erecta, hasta ca. 40 cm alt, glabrescente. Tallo 
solitario, laxamente ramificado desde la base o mas arriba, poco hojoso. Hojas 
sésiles, las basales escasas, con lamina espatulada, base atenuada en 
pseudopeciolo y apice agudo, hojas caulinares sésiles, ovadas a anchamente 
ovadas, agudas, con 3-5-nervios sobresalientes en el hipofilo, de 10-44 mm long. 
x 5-15 mm lat.,con tricomas glandulares no capitados en la base foliar adaxial. 
Flores terminales solitarias 0 en cimas 3-5-floras, raro 8-floras y flores axilares 
solitarias o geminadas; pedicelos de 20-60 mm long.; caliz acrescente y 
persistente, de 10-16 mm long., lobulos angosto-ovados, agudos, desiguales, hasta 
3 veces mas largos que el tubo, tubo con anillo de tricomas glandulares 
pluricelulares en su borde inferior interno, de 3-5 mm long. corola lila, violacea 
oscura en la base de cada lobulo, de 17-26 mm long., tubo breve (5-7 mm long.), 
glabrescente, con tricomas glandulares pluricelulares por encima de la union 
de los estambres, Lobulos elipticos a espatulados, agudos, de 10-19 mm long., 3- 
5 veces mas largos que el tubo, nectarios epipétalos en U; filamentos estaminales 
planos, mas largos que la altura media de la corola e insertos cerca de la mitad 
inferior del tubo en las flores perfectas, anteras versatiles (primero introrsas, y 
extrorsas en la dehiscencia), elipsoides, inclusas, de + 3 mm long, tecas libres en 
su tercio inferior; en las flores pistiladas, estambres reducidos a estaminodios 
(filamentos breves con anteras atrofiadas, vacias); gineceo angosto-elipsoide, 
sésil, lobulos estigmaticos ovados. Capsula de color marron claro, cuando 
madura excede la corola persistente. Semillas numerosas, castaho oscuras, 
subesféricas, menores de | mm de diam., con tegumento levemente reticulado. 

Etimologia.—El epiteto se debe a Humberto A. Fabris 1924-1976), destacado 
docente e investigador botanico argentino, quien colecciono6 el material tipo y 
dedico una buena parte de su vida al estudio de las Gentianaceae, en especial 
Gentianella, de América del Sur. 

Observacion.—En la coleccién tipo de G. fabrisii se encontraron semillas 
maduras provenientes de frutos derivados tanto de las flores pistiladas como 
de las perfectas (Fig. 1 F-H). 

Habitat y distribucién geografica.—Hasta el momento, solo se conoce esta 
especie por el material tipo, proveniente de las montanas del oeste de la Prov. de 
Tucuman, por arriba de los 3000 m. Al igual que sus congéneres, habita en 
praderas humedas de altura donde convive con Gentianella bromifolia (Griseb.) 
T.N. Ho et S.W. Liu. 


FILIPPA AND RARRNZA 


AMERICA DEI Sl 


JR 


Fic.1 tianella fabrisii.A. Habito. B. Corolad 


1. do fl re | 


I 
E. Caliz, vista externa. F. Embrion.G. Semilla, vista 
erie ore sr. Lb la . ime 1 


1 

y 
lateral. H.T 
Py I PY -l 


K. Flor, vista lateral. L. Botén floral despl 10, ot 
fi I ' 


E, I,K, L. De Fabris 14174. 


BRIT.ORG/SIDA 22(1) 


Gentianella fabrisii es afin a Gentianella bromifolia por compartir la 
ramificacion de los tallos, la longitud de los peciolos, la disposicion de las flores, 
y la forma y el color de los lobulos corolinos; se diferencian por su sexualidad 
(planta ginomonoica vs. planta monoico-monoclina), la forma de las hojas 
caulinares (ovadas a anchamente ovadas vs. linear-elipticas), la forma de los 
lobulos calicinos (angosto-ovados, agudos vs. linear-triangulares, aleznados), 
la relacion longitud tubo/Id6bulos de la corola (1/3-1/5 vs. 1/2-1/3) y por el 
gineceo (sésil vs. estipitado). 

En los tratamientos de Gilg (1906, 1916), ya se hace mencion de la presencia 
de poligamia entre las especies de Gentianella (sub nom. Gentiana). Si bien la 
mayoria de las especies poseen flores monoclinas, se pudo comprobar que de 
las 29 especies que crecen en la Argentina, en 5 de ellas ocurre ginomonoecia, 
entre las que se encuentra G. fabrisii,en tanto que solamente G. florida presenta 
ginodioecia Juarez de Varela 1997; Filippa & Barboza, inédito). Un caso muy 
particular es el de G. thiosphaera debido a la ocurrencia de trimonoecia (segun 
Font Quer 1979), es decir flores perfectas en su mayoria, y pocas estaminadas y 
pistiladas en un mismo individuo (Filippa & Barboza 2001) 


LAS 7 ESPECIES POLIGAMAS DE LA ARGENTINA 
SE DIFERENCIAN EN LA SIGUIENTE CLAVE 


pe cee terminales densas. Anteras hasta de 2 mm long. Pedicelos cortos, 
en oe o blanco-amarillenta, con tubo notablemente mas corto que los 
lObulos. Anteras completamente hee Plantas aes G. florida 


Corola amarillao lila cina,cont {ny j {| it { 


Anteras inclusas 0 apenas exertas, Plantas no pines ee 
37 L I: 4 eet | IAtaral 


es ae mm. ). Plantas trimonoicas G. thiosphaera 
1 | 


3. 4s largo que los ldbulos. Gineceo sésil. Plantas 
ginomonoicas. 
4, ils amarilla. Flores axilares en verticilos. Hojas opuestas o en verticilos 
G. multiflora 
4 ae ere Flores axilares no verticiladas. Hojas siempre opuestas 
G.imberbis 
1. Infl i inales | bien fl lit Anteras de 2-4 mm.Pedicelas 
comparativamente largos, de 15-90 mm 
5. Tallo solitario. Hojas ovadas a anchamente ovadas. Lobulos calicinos angasto- 
vados G. fabrisii 
5. Tallos numerosos. Hojas elfpticas 0 angosto- a linear-elipticas. Lobulos calicinos 
ara a linear-triangulares. 
6. Hojas caulinares elipticas, siem} puestas. Nectarios en V. Semillas de 1-1,1 
mm X 0,7-0,9 mm G. kurtzii 


. Hojas caulinares angosto- a linear-elipticas, opuestas o en verticilos trimeros. 
Rlactarias en U. Semillas de 0,5-0,9 mm X 0,3-0,4 m 


G. myriantha 


CILIDDA AND RARRNZA AMERICA NEI SUR 133 


NUEVOS SINONIMOS PARA ESPECIES DE GENTIANACEAE 
CICENDIA 


Cicendia quadrangularis (Lam.) Griseb., Gen. sp. Gent. 157. 1839. Basonimo: Genti- 
uadrangularis Lam., Encycl. 2:645. 1788. Tipo: PERU: “Environs of Lima,” Dombey s.n. 
(HOLOTIPO: P!) 


Gentiana flava Larranaga, Escritos de Don D.A. Larranaga 2:100. 1923. Tipo: No citado. Syn. nov. 


Si bien el herbario de Larranaga fue destruido por accion de los insectos (Alonso 
Paz, com. pers.), gran parte de su obra escrita pudo rescatarse y fue publicada en 
varias etapas; en 1923, se dieron a conocer las descripciones de las especies y 
cuatro afios mas tarde (1927) - en el Atlas (Parte I. Botanica)-, las magnificas 
ilustraciones de referencia. 

Cuando Larranaga describe Gentiana flava no precisa ningun material tipo, 
solo sefiala que se encuentra “..en todos los prados; pero es tan pequena que se 
confunde muy facilmente con el pasto tierno entre que se oculta.” La diagnosis 
es breve (planta muy pequena, tallo poco ramificado y unifloro, hojas lineares, 
corola tetramera, glabra, amarilla, hipocrateriforme); sin embargo, los caracteres 
que utiliza, refrendados por la ilustracion (Lam. LXXV1J), son lo suficientemente 
precisos para decidir su sinonimia con Cicendia quadrangularis. 


CURTIA 


Curtia tenuifolia Ce oe Bot. Zentralbl. 60:357. 1894. (Fig. 2). Basonimo: 
Hist - Guiane 1:70, .26, f.2. 1775. Tipo: “Habitant in pratis humidis 


Ca z & Guian ae’ ‘ | 

lensis Ar , Anales Mus. Nac id 2, 1:65, f.3. 1911. Tipo: [URUGUAY] 
“Montevideo, arenales hamedos de Carrasco,” Abr 1874, J. Arechavaleta 163 (HOLOTIPO: MVM)). 
Syn. nov. 


Arechavaleta (1911) describe C. montevidensis sin mencionar los caracteres que 
la diferencian de sus congé El habito y la disposicion, forma y longitud de 
las hojas (opuesta, linear-subulada y ca. 4 mm long.) y la longitud del caliz y 
corola (ca.4 mm _y 5mm long. respectivamente) observados en el material tipo, 
son coincidentes con los de C. tenuifolia. 

Por otro parte, cuando Fabris (1973) cita por primera vez la existencia de 
Curtia malmeana en la Argentina, destaca su afinidad y posible sinonimia con 
C. tenuifolia (esta situacion esta propuesta en una preliminar Checklist de Bo- 
livia virtual: www.efloras.org). Mediante el analisis del material tipo de C. 
malmeana (Brasil: Malme 424b, S!), corroboramos esta sinonimia. De esta 
manera, Curtia tenuifolia (Fig. 2), es la Gnica especie que crece en la Argentina 
(Prov. Corrientes) y sur de Uruguay (Dpto. Montevideo y Canelones). 


GENTIANA 


Gentiana prostrata Haenke, Collectanea 2:66, pl.17, f.2. 1789. Tipo: ‘Rara habitat in 
jugis nivosis alpium Kartal et Frosnitz, principatus Salisburgensis...”. (n.v.). 


134 BRIT.ORG/SIDA 22(1) 


Fic. 2. Curtia tenuifolia. A. Habito. B. Corola despelgada. C. Gineceo. D—-F. Anteras en vista ventral, dorsal y lateral 


respectivamente G. Semill lado. H. Flor. I. F 


CILIDDA AND RADRNZTA AMERICA NEI SUR 135 


Gentiana minima Phil. Linnaea 29:13. 1858. Tipo: CHILE: “In Andibus interioribus prov. Santiago,” 
Philippi s.n. (HOLOTIPO: SGO 053950). Syn. nov. 
Fabris (1953) hace referencia al gran polimorfismo que presenta G. prostrata, lo 
que ha motivado una profusa sinonimia para esta especie (Fabris 1983). Genti- 
ana minima, descripta para Chile, ha pasado inadvertida en la literatura de los 
ultimos anos. El analisis del holotipo depositado en SGO, puso en evidencia que 
se trata de un sindnimo mas de G. prostrata. 


GENTIANELLA 

Gentianella cosmantha (Griseb.) J.S. Pringle, Phytologia 48:281. 1981. Basonimo: 
Gentiana cosmantha Griseb., Pl. lorentz.: 161. 1874. Tipo: ARGENTINA. TUCUMAN: Cuesta de 
la Casilla, Apr 1872, Lorentz 774 (HOLOTIPO: Bl; ISOTIPO: CORD)). 


Gentianella cosmantha f. violacea Fabris, en A.L. Cabrera (dir), Flora prov. Jujuy 8:66. 1983. Tipo: 
ARGENTINA. Jujuy: Dpto. Capital, Laguna de Yala, 26 Mar 1979, A.L. Cabrera et al. 30678 
(HOLOTIPO: SI). Syn. nov. 


Fabris (1983) dif ia laf. violacea de la f. cosmantha por el tamano y color de 
la corola (mas pequena y de color violaceo en vez de rojo 0 rojo-anaranajado). 
Las observaciones realizadas a campo y de ejemplares de herbarios provenientes 
del area de distribucion de Gentianella cosmantha, revelaron que poblaciones 
con corolas rojizas poseen tamano dentro del rango de variacion senalado por 
Fabris para la f. violacea. Con respecto al color, es frecuente que al envejecer la 
flor, la corola vire de rojo a rojo-vinoso. 


Gentianella magellanica (Gaudich.) Fabris ex D. M. Moore, Sci. Rep. Brit. Antarc. 
Surv. 60:103. 1968. BAsonimo: Gentiana magellanica Gaudich., Ann. Sci. Nat. (Paris) 
seas 102. 1825. Tipo: [ARGENTINA| “Iles Malouines,” Gaudichaud s.n. (HOLOTIPO: B+; ISOTIPOS: 


Gentiana patagonica & darwinii Griseb., Gen. sp. Gent. 237. 1839. Tipo: CHILE. “Ad fretum 
Magellanicum, Elizabeth Island,” Darwin s.n. (HOLOTIPO: K!). Syn. 
ti Phil., Linnaea 29:109. 1858. Tipo: [CHILE]: “In ae Bee dictis ad altit., ca. 
ed.s.m.,3 ie 1837,” F Fonck 130 SGO 0539206!). Syn. nov. 
eee lactea Phil., Anales ce eae 90: 206, 1895, Tipo: [CHILE]. VALDIVIA: “Cordillera Pelada, 
1889,” F Philippi s.n. (HOLOTIPO: SGO aon ISOTIPO: BM!). Syn. nov. 


Gentiana patagonica var. gracilis v, Revista Mus. La Plata, Secc. Bot. 7:348. 
1896. TIPO: [ARGENTINA “Vallee de POlivaia, cr ea 1896,” N. Albov 661 y 662 (SINTIPOS: 
LP!). Syn. nov. 


Gentiana pearcei f. nana Hauman, Anales Soc. Ci. Argent. 86(3-4):299. 1918. SinTIPOs: [ARGEN- 
ROV. MENDOZA|: “Las Cuevas, Sep-Oct 1909,” M.S. Pennington 20 (BAF); “Cordillére de 
Tunuyan, 4 2700 metres,” leg. Sanzin 1301 (Herb. desconocido). Syn. nov. 

Los caracteres utilizados por Grisebach para Gentiana patagonica { darwinii 
(numero de piezas de la corola) asi como los empleados por Philippi para Gen- 
tiana lactea (grado de ramificacion de los tallos, forma de las hojas e 
inflorescencias paucifloras) y los usados por Philippi, Albov y Hauman para 
Gentiana modesta, Gentiana patagonica var. gracilis f. pumila uniflora y Genti- 


136 BRIT.ORG/SIDA 22(1) 


ana pearcei f. nana respectivamente (altura de la planta y numero de flores), 
coinciden en un todo con los que definen a Gentianella magellanica 

Por otro lado, Spegazzini (1902) describio algunos taxones infravarietales de 
Gentiana magellanica(G. magellanica var. darwinti f. cyanescens, G. magellanica 
var. darwinii f. albiflora; G. magellanica var. typica f. cyanescens; G. magellanica 
var. typica f. albiflora) que coinciden con el rango de variacion del color de la 
corola de Gentianella magellanica. Estos nombres subespecificos carecen de toda 
validez taxonomica, ya que no se mencionan especimenes tipos ni ilustraciones. 


Gentianella multicaulis (Gillies ex Griseb.) Fabris, Bol. Soc. Argent. Bot. 7:89. 1958. 
BASONIMO: Gentiana multicaulis Gillies ex Griseb., Gen. sp. Gent. 225. 1839. Tipo: [ARGEN- 
TINA. MENDOZAl “Habitat in humidis pr. Uspallata et pr. Manantiales del Portillo in Andibus 
Mendozianis, Fl. Jun.,” J. Gillies s.n. GHOLOTIPO: K n.v.). 

Gentianella achalensis (Gilg) T. N. Ho et S. W. Liu, Bull. Brit. Mus. (Nat. Hist.), Bot. 23(2):61. 1993. 
BASONIMO: Gentiana achalensis Hieron, ex Gilg, Bot. Jahrb. Syst. 22:322. 1896. Tipo: ARGEN- 
TINA, CORDOBA: San Miguel, en la Sierra Grande (Achala), 27 Mar 1875, J.H.E.W. Hieronymus 
215 (LECTOTIPO: CORD!)). Syn. nov. 

Gentianella mendocina (Gilg) J. S. Pringle, Sida 1368. 1986. BASONIMO: Gentiana mendocina Gilg, 
Bot. Jahrb. Syst. 54, 2 (Beibl. 118):36. 1916. Tipo: ARGENTINA. MENDOZA: 1876, R.A. Philippi s.n. 
(HOLOTIPO: B+). Syn. noy 


OC. ( 


Al igual que G. magellanica, G. multicaulis es una especie muy variable tanto en 
sus caracteres vegetativos como florales. El rango de variacion se observa en el 
numero, grosor y altura de los tallos, en la presencia y cantidad de hojas basales, 
en la cantidad de oe Bas ya ae cencia y en el color de la corola. El lectotipo 


hal 


de Gentianad ac Gentianella multicaulis. De igual 


manera, la diagnosis de G. mendocina se ajusta a dicha especie. 

Gentianella ottonis (Phil.) Munoz, Sinopsis Fl. Chile: 134. 1959. Basonimo: Gentiana 
ottonis Phil. Linnaea 33:177. 1805. SINTIPOS: ARGENTINA. MENDOZA: “Portillo de los Puiquenes,” 
R.A. Philippi s.n. (B+); CHILE: “Andes de Santiago,” R.A. Philippi s.n.(SGO 053960). 


iana ottonis var. hexamera (Phil.) Reiche, Fl. Chile 5:131.1907. BASONIMO: Gent hexamera 
Phil, Anales Univ. Chile 43:510. 1873. Tipo: CHILE: “Valle del Yeso de la Cordillera de San- 
tiago, Ene 1861,” R.A. Philippi s.n. (HOLOTIPO: SGO 053939!, 041974! ISOTIPO: BAF!). Syn. nov. 


Gen 


Cuando Reiche (1907) subordina G. hexamera a nivel varietal de G. ottonis, 
senala que los caracteres de esta especie (tallos unifloros y flores 6-7-meras) 
podrian corresponder a un hecho tanto anémalo como frecuente. Si bien la 
mayoria de los ejemplares analizados de G. ottonis presentan flores pentameras, 
las plantas de las dos cartulinas del holotipo y la del isotipo de G. hexamera, no 
dejan dudas de esta sinonimia por su habito rizomatoso, el tamano de las flores 
y la forma de los l6bulos corolinos. 


ZYGOSTIGMA 


Zygostigma australe (Cham. & Schltdl.) Griseb., Gen. sp. Gent. 150. 1839. Basonimo: 
Sabatia australe Cham. & Schltdl, Linnaea 1:194. 1826. Tipo: BRASIL: “Prov. Cisplatina, 
Braxuelo, Novembri mense floret.,” Sellow s.n. (W?). 


Gentiana rubra Larrafaga, Escritos de Don D.A. Larranaga 2:100. 1923. Tivo: No citado. Syn. nov. 


FILIPPA AND RARRO7ZA AMERICA DEL SUR 137 


Tal como en el caso de Gentiana flava, Larranaga describe e ilustra esta especie 
con bastante precision, sin mencionar ejemplar de referencia, salvo su nombre 
vulgar y fecha de coleccion: “Canchalagua, Ener. 1811.” En base al protdlogo, no 
quedan dudas que se trata de Zygostigma austra 


© 


Sobre la ampliacion del area de distribucion de Chelonanthus viridiflorus en 
América del sur. 

Chelonanthus (Griseb.) Gilg es el género neotropical con mayor area de 
distribucion de la tribu Helieae Gilg, extendiéndose desde el norte de México 
hasta el sur de Brasil y Bolivia; hasta el t eptanca.8 (Struwe 
et al. 2002). Su taxonomia es muy confusa; en efecto, muchas ae sus especies 
han sido descriptas o incluidas en distintos géneros tales como Lisyanthus 
Aublet (Don 1838; Grisebach 1839, 1845; Progel 1865, Bentham 1876), Helia Mart. 
(Kuntze 1891) e Irlbachia Mart. (Maas 1985; Pringle 1995). En la actualidad, 
algunas especies transferidas a Irlbachia (Maas 1985), han sido rehabilitadas a 
Chelonanthus (Struwe et al. 1998, 1999) por lo que su circunscripcion no resulta 
facil. Por otro lado, al no haber un tratamiento monografico del género atn 
permanecen sin resolverse muchos problemas nomenclaturales y de 
circunscripcion de taxones (Struwe et al. 2002). 

En la Argentina, y como consecuencia de los cambios en la taxonomia de 
Chelonanthus, el registro de sus especies ha sido variable. En 1953, Fabris 
identifico como C. grandiflorus (Aubl.) Hassl., un Gnico ejemplar proveniente 
del litoral (Prov. Misiones: Molfino s.n.). Hunziker 1984) puso en duda la 
existencia de esta especie en ese pais debido a su escasa representatividad en 
los herbarios argentinos. Trabajos recientes registran la ocurrencia de Irlbachia 
alata (Aubl.) Maas en estudios p de especies del litoral fluvial (Pire et al. 
2004) e L alata subsp. viridiflora (Mart.) Pers. @ Maas en estudios floristicos de 
los esteros del Ibera, en Corrientes (Arbo & Tressens 2000). Al analizar los 
ejemplares de referencia y otras colecciones del nordeste de la Argentina, se pudo 
verificar que solo Chelonanthus viridiflorus habita en este pais; asi, su area de 
distribucion se extiende hacia el sur del continente. 

En esta contribucion se proporciona una detallada descripcion y se ilustra 
C. viridiflorus a la par que se adicionan otras colecciones que reafirman la 
distribucion de esta especie en territorio argentino y paraguayo. 


1c 


1. Chelonanthus viridiflorus (Mart.) Gilg, en A. Engler u. K. ee Nat. 
Pflanzenfam. 4(2):98. 1895. (Fig. 3). Lisianthus viridiflorus Mart., Nov. 1. 
2(2):94, t.173. 1827. Helia viridifl Mart.) Kuntze, Revis. gen. pl. 2:428. 1891. hie oe 
subsp. eel (Mart.) oe & Maas, Proc. Kon. Ned. Akad. Wetensch., C 88(4):410. 1985. 
Tipo: “Crescit in subalpinis campis prope Villa Rica, Sabara, et alibi in Provincia Minarum” 
Martius 72 (SINTIPO: MI). 


Hierba anual. Tallo erecto, simple, terete en la base y subcuadrangular 
superiormente, de 65-100 cm alt. Hojas enteras, pinnatinervias, las inferiores 


138 


BRIT.ORG/SIDA 22(1 


hel fiflorus. A. Habito. B. Semilla. C. Corola desplegada con lusos. D. Transcorte de 
ovario. E. Antera. F. Frut aliz y estilo persistentes. G. Frut dehi ia. H. Flor. 1. Tri glandular del caliz.J 
Gineceo. K. Caliz, vist 1o las glandulas.L. pical de infl A, E-G de Schinini et al.11248; 
B de Arbo etal. 8492; C, D, H—L de Molfino s.n. La escala de 12 mm vale para C,J,L;lad 
FH. 


FG K lad 


CIIDDA AND RAPRNTA AMERICA NEI SUR 139 


brevemente pecioladas, subconnadas (las bases confluyen en una vaina 
membranosa), las superiores sésiles, lamina eliptica, aguda, base atenuada, de 
(1,5-)2,5-9 cm de long. x (0,6-)1,5-4,5 cm de lat., hipofilo con 2 6 3 pares de 
nervios secundarios, opuestos y sobresalientes, peciolo de 4+,5-6 mm de long. 
Cimas dicotomicas terminales, por lo general paucifloras, cada rama con 2-6 
flores, bracteas elipticas, agudas. Flores pentameras, perfectas, levemente 
zigomorfas, horizontales, reflexas en post-antesis, cortamente pediceladas (5-7 
mm); bractéolas escuamiformes, hasta ca.7 mm de long, caliz campanulado, 
persistente y poco acrescente, de 6-9 mm de long., tubo muy breve, con tricomas 
glandulares pluricelulares en la base, de 2-3 mm, l6bulos crasos, ovados, obtusos, 
por fuera y en el centro un area glandular elipsoidea sobresaliente; margen 
translucido; corola verde palida, de (2-)2,5-5 cm de long,, tubo angosto en la 
base, ensanchado en campana ventricosa por arriba del caliz, de (1,6-)2-4,4 cm 
de long., lobulos ovados, rotundos, pentanervados, hasta unas 7 veces mas cortos 
que el tubo, apice grueso y oscuro, recurvado; estambres insertos debajo del 1/3 
inferior del tubo, filamentos laminares subiguales, anteras ovoides 0 sagitadas, 
libres en su 1/3 basal, con un apéndice estéril en el apice, polen en tétrades, 
blanco, abundante; gineceo bicarpelar, ovario claviforme, placentas introflexas, 
estilo filiforme, de igual longitud que los estambres, en post-antesis igual o mas 
largo que la corola, estigma en dos lamelas ovadas. Capsula septicida, reflexa, 
ovoide-cénica, rostrada (estilo persistente), dehiscente por el medio, en dos 
valvas al madurar. Semillas numerosas por fruto, anguladas, menores de 1 mm 
de diam. 

Habitat y distribucion geogrdfica.—Especie caracteristica de zonas 
pantanosas, sabanas htimedas, borde de rios y embalsados. Habita en Venezu- 
ela, Colombia, Guyana, este de Bolivia (Dpto. Santa Cruz), centro y sudeste de 
Brasil (Edos. Mato Grosso, Goias, Sao Paulo), sudeste de Paraguay (Dptos. Alto 
Parana, Caaguazu, Canendiyt, Itapua y San Pedro) y noreste de Argentina (Provs. 
Corrientes y Misiones). 


Material examinado: ARGENTINA. Prov. Corrientes. Dpto. Ituzaingo: Desembocadura del Arroyo 
Garapé en el Rio Parana, 45 km al E de Ituzaing6, en pantano, 24 Abr 1975, Schinini et al. 11248 (CTES), 

Dpto. Santo Tomé: Reserva Natural Provincial del Ibera, Laguna Galarza, 28° 05'S, 56° 40' W, 0-100 
m,embalsado al borde del canal de a ala Laguna, 25 Abr 1995, Arbo et al. 6553 (CTES), Laguna 
de Luna, 28° 04'S, 56° 49’ W, costa occidental, en embalsado, 24 Nov 1999, Arbo et al. 8492 (CTES); 
Laguna Galarza y Luna, 03 Abr 1998, Honfi & Davina 00895 (MNES); Laguna Galarza, embalsado, 9 
Dic 1974, Quarin et al. 2509 (CTES). Prov. Misiones. Dpto. San Javier: San Javier, Feb 1922, Molfinos.n. 
ae PARAGUAY. Dpto. Caaguazu: Yhu, 24 Ene 1951, Sparre et Vervoorst 2060 (CTES, LIL); 10 km al 


e Caaguazu, camino a Yhu, orilla de arroyo en terreno pan 0, 8 Feb 1966, Krapovickas et 
i 9H ok Dpto. Canendiyu: Car pa-cué, pastizal sobre slo sees se Mar 1996, Jimenez & 
Marin 110 (CTES, PY); Ballinoti cué, cerrado, eas al b turad Feb 1988, Jimenez & 
eae BJ (CTES). Dpto. Itapua: Isla Yacyret, f I que rod ee de Arary, 22 Abr 1993, 


Quintana et al. 265 (CTES, PY). 


140 BRIT.ORG/SIDA 22(1) 


Sobre la presencia de Schultesia guianensis en Argentina y Paraguay 

La cita de Fabris (1953) para Schultesia guidnensis en la Argentina estaba 
documentada solo por un ejemplar del Prof. J. Molfino s.n., procedente de 
Misiones. Se pudo localizar este ejemplar en el herbario BAF y corroborar que, 
efectivamente se trata de esta especie al compararlo con el material tipo 
depositado en Paris. En este trabajo, se describe e ilustra a esta especie y se la 
documenta ademas, para Paraguay. 

Schultesia guianensis (Aubl.) Malme, Ark. Bot. 30.2):9. 1904. (Fig. 4). Exacum 

eguianensis Aubl, Hist. pl. Guiane 1:68, t.26, f.1. 1775. Tipo: “Guiane” (PD). 


Hierbas anuales, pequenas, glabras, de 4-15 cm alt. Tallo erecto, tetragono, con 
alas reducidas en los angulos, poco ramificado superiormente. Hojas enteras, 


opuestas, las inferiores pseudopecioladas, espatuladas, obtusas, de 7-10 mm de 
long. x 5mm de lat.; las superiores sésiles, elipticas, agudas, subconnadas en la 
base, de 10-25 mm de long. X 5-8 mm de lat. Flores solitarias 0 en cimas 
dicotomicas en el apice de las ramas; bracteas elipticas, agudas. Flores 
tetrameras, perfectas, sésiles o con pedicelos breves (hasta de 5 mm); caliz 
urceolado, persistente, poco acrescente, de 12-19 mm de long., 4-alado, alas 
marcadamente retinervadas (1,5-2 mm de ancho), tubo de 8-11 mm, l6bulos 
subtriangulares largamente aleznados; corola rosada a purptrea, marcescente, 
e 15-22 mm de long., tubo angosto en la garganta, de 9-12 mm, lobulos 

obovados, rotundos, a veces apiculados, pentanervados, un poco mas cortos que 
el tubo; estambres insertos en la 1/2 basal del tubo corolino, inclusos 0 apenas 
exertos, subiguales, sector libre de los filamentos alado hasta su altura media, 
alas irregularmente partidas en el apice; anteras ovoides 0 sagitadas, libres en 
su 1/3 basal; polen en tétrades; gineceo bicarpelar, ovario elipsoide, un poco mas 
corto que el tubo corolino, placentas introflexas, estilo filiforme, deciduo, estigma 
con dos lamelas trans-ovadas a subcirculares. Capsula septicida, elipsoide. 
Semillas numerosas por fruto, subesféricas, ca | mm de didm.; episperma 
reticulado, células poligonales con paredes ligeramente onduladas. 

Observacion.—Acorde con Struwe et al. (1999) y Pringle (com. pers.), los 
especimenes procedentes de otras areas de América muestran algunas 
diferencias con respecto a nuestros datos, sobre todo en la altura de la planta y 
la longitud del caliz y la corola. Asi, Schultesia guianensis en su conjunto, 
presenta un rango de variacidn mas amplio en cuanto a los caracteres 
mencionados, extendiéndose la altura de la planta desde 3-45 cm, la longitud 
del caliz de 8-22 mm y, la de la corola de 8-26 mm 

Habitat y distribucion geogrdfica.—Especie palustre de amplia distribucion 
tropical y subtropical, que se extiende desde México hasta el noroeste de Ar- 
gentina y Paraguay. En el Cono Sur habita en campos bajos y anegados en la 
ribera de cursos de agua y lagunas. 


FILIPPA AND RARRNZA AMERICA DEL SUR 141 


& 
Fic. 4. Schultesi j is. A. Habito. B. Caliz abierto.C. Corola { y gi D. Semilla | i f ticul 
De Molfino s.n. (BA 52976). lade5 le para B, C 


142 BRIT.ORG/SIDA 22(1) 


Material examinado: ARGENTINA. Prov. Misiones. Dpto. Capital: Posadas a San José, Feb 1922, Molfino 
s.n(BA 52976); Posadas, hacia San José, en campo bajo, Molfino s.n., Feb 1922, (BAF). PARAGUAY. Dpto. 
Cordillera: In regione lacus rte: 1913, Hassler 12596 (BAF, LP); Camino de Altos a Loma Grande 
25° 17'S, 55° 15' O, 20 Mar 1989, N. Soria 3438 (FCQ); Cordillera, 25.08° S 57.12' W, 25 Febr 1990, E. 
Zardini y R. Velazquez awe (AS). BRASIL. Edo. Rio de Janeiro: Cabo Frio, Oct 1899, E. Ule 4747 
(CORD). BOLIVIA. Dpto. Santa Cruz: Fortin Ra-velo-Cerro San Miguel, Mar 1989, F. Mereles 2744 
(PY). 


AGRADECIMIENTOS 


Las autoras expresan su gratitud a los Dres. JS. ae y E.F Guimaraes por la 
lectura critica de este trabajo y sus acertadas observaciones para mejorar el 
manuscrito. A los curadores y asistentes curatoriales de los herbarios AS, BA, 
BAP BM € CORD CLES FCO. KLE LP VMN ES MYNME Pi 5, 5OU. SL yy, 
Z, quienes pusieron a nuestra disposicion el material de herbario estudiado. Al 
Dr. P. Maas por habernos facilitado la obtencién de bibliografia especifica. A M. 
Ehn (S) y a los Dres. M. Mufioz (SGO) y H.S. Osorio (MVM) por el envio de 
imagenes digitales de materiales tipo. Al Dr. E. Alonso Paz (MVFQ) por la 
informacion brindada sobre el herbario de Don D. Larranaga. Al Dr. L. Ariza 
Espinar por suconstante estimulo. A los dibujantes L. Ribulgo y P. Peralta por la 
confeccion de las ilustraciones. Al Consejo Nacional de Investigaciones 
Cientificas y Técnicas (CONICET Argentina), Agencia Cordoba Ciencia (ACC- 
Argentina), Secretaria de Ciencia y Tecnologia (UNC-Argentina) y a la Myndel 
Botanica Foundation (Argentina) por el apoyo financiero otorgado. 


REFERENCIAS 


Arso, M.M. & S.G. Tressens (eds.) 2000. Flora del Ibera. EUDENE, Corrientes. Pp. 1-613. 

ARECHAVALETA, J. 1911. Vegetacion Uruguaya. Varias especies nuevas. Anales Mus. Nac. 
Montevideo, ser. 2, 1:59-83. 

BentHAM, G. 1876.Gentianeae. En:G. Bentham & J.D. Hooker,eds.Genera plantarum 2. Lovell 
Reeve & Co.,Williams & Norgate, London. Pp. 799-820. 

Don, G. 1838. A general history of the dichlamydeous plants 4. J.G.& F. Rivington et al. 
London. Pp. 173-214 

Fasris, H.A. 1953. Sinopsis preliminar de Gentiandceas argentinas. Bol. Soc. Argent. Bot. 
4:233-259. 

Fasris, H.A.1973.Un nuevo género de Genciandceas para la Flora Argentina, Curtia Cham. 
& Schidtl. Bol. Soc. Argent. Bot. 15:130-131. 

Fasris, H.A. 1983. Gentianaceae. En: A.L. Cabrera, ed., Fl. Prov. Jujuy 8. Coleccion Cientifica 
Instituto Nacional de Tecnologia Agropecuaria, Buenos Aires. Pp. 55-85 

Fivippa, E. & G.E. Barsoza. 2001. Novedades en Gentianaceae de Argentina. Kurtziana 29: 
79-83. 

Fiuippa, EM & G.E. Barsoza. 2003. Novedades en Gentianaceae de Argentina. Il. Kurtziana 

65-67 
Fiippa, E.M & G.E. Barsoza. Gentianaceae, Flora Fanerogamica Argentina. PROFLORA- 


FILIPPA AND RARRNTA AMERICA NEI SUR 143 


CONICET. Inédito. 

Font Quer, P. 1979. Diccionario de Botanica. Ed. Labor, S.A. Barcelona. Pp. 1-1244. 

Gi, E. 1906. Beitrage zur Kenntnis der Gentianaceae Ill. Gentianaceae andinae. Repert. 
Spec. Nov. Regni Veg. 16-17:33-56. 

Gitc, E. 1916. Gentianaceae andinae. Monographischem Zusammenstellung der Genti- 
ana-Arten Stid-Amerikas. Bot. Jahrb. Syst. 54-2. Beibl.118:80-85. 

GRrISEBACH, H.R. 1839. Genera et species Gentianearum.J.G. Cotta, Stuttgart & Tubingen. Pp. 

364 


GriseBACH, H.R. 1845. Gentianaceae. En: A. De Candolle, ed. Prodromus systematis naturalis 


regni vegetabilis 9. Fortin, Masson et Sociorum, Paris. Pp. 38-141. 

Hunziker, A.T. 1984.Los géneros de Fanerdgamas de Argentina.Claves para su identificacion. 
Bol. Soc. Argent. Bot. 23:158 

JENSEN, S.R. & J. ScHripsema. 2002. Ct t and pharmacology of Gentianaceae. En: 
L. Struwe and V.A. Albert, eds. Gentianaceae - Systematics and Natural History. Cam- 
bridge University Press, Cambridge. Pp. 573-631. 

JUAREZ DE VARELA, F. 1997. Gentianaceae. En: Flora del Valle de Lerma, Aportes Botanicos de 
Salta, Ser. Flora 4:1-19. 

Kuntze, C.E.O. 1891.Revisio generum plantarum 2. Arthur Félix, Leipzig. Pp. 426-432. 

Larraniaca, D.A. 1923. Escritos de Don Damaso Antonio Larrafaga. Il. Botanica-Zoologia. 
Instituto Histdrico y Geografico del Uruguay, Montevideo. Pp. 1-512. 

LARRANAGA, D.A. 1927. Escritos de Don Damaso Antonio Larranaga. Atlas, Parte |. Botanica, 
Instituto Histérico y Geografico del Uruguay, Montevideo. Lam I-CXXxXI. 

Maas, PJ.M.1985.Nomenclatural notes on Neotropical Lisyantheae (Gentianaceae). Proc. 
Kon. Ned. Akad. Wetensch C 88:405-41 2. 

Prinate, J.S.1995.Gentianaceae. En:G. Harling and L. Anderson, eds. Flora of Ecuador 53.Pp. 
Sie 

Procet, A. 1865. Gentianaceae. En: C.F.P von Martius, ed. Flora Brasiliensis 6 (1). Leipzig. Pp. 
197-248. 

Pire, S.M., LM. AnzoteGu! & G.A. Cuanrano. 2004. Estudios palinoldgicos en el Litoral fluvial 
argentino. Instituto Superior de Correlaci6n Geoldgica, Ser. Misc. 12:139-146. 

ReicHe, C. 1907. Jencianaceas, Fl. Chile 5 [Gentianaceae: 120-134]. 

Specazzini, C. 1902. Nova addenda ad Floram Patagonicam Il. Anales Soc. Ci. Argent. 53 
[Gentianaceae: 73-75]. 

Stauwe, L.& V.A. Aceert. 1998. Lisianthus (Gentianaceae), its probable homonym Lisyanthus, 
and the priority of Helia over Iribachia as its substitute. Harvard Pap. Bot. 3:63-71. 

Struwe, L., PJ.M. Maas, O. PIHLAR & V.A. AcperT. 1999. Gentianaceae. En: PE. Berry, K. Yatskievych 
& B.K. Holst, eds. Fl. Venezuelan Guyana 5:474-542. 

Struwe, L., J.W. Kavereit, J. KLACKENBERG, S. Nisson, M. THiv, K.B. Von HAGEN & V.A. AvBerT. 2002. 
Systematics, character evolution, and biogeography of Gentianaceae, including a new 
tribal and subtribal classification. En:L. Struwe and V.A. Albert, eds. Gentianaceae - Sys- 
tematics and Natural History. Cambridge University Press, Cambridge. Pp. 21-309. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Oi Lers. 2005?. Coffee, cappuccino, espresso and much more. (ISBN 3-921799- 
48-1, hbk.). Kochbuch-Verlag?, Munich. $30.00, 159 pp, 14 color illus., 6 3/4" 


x 8 3/8". 


Robusta, arabica, peaberry, mocha...? 
“Perfectly home made by Olli Leeb,” follows the title of this charming homespun publication. 
A loosely edited translation from the German results in a quaint posualys across-the-table Bo of 
} 
ur i€ if 


narrative. Add to that the ty pevnile: styled, unjustified, co 
fun book! And asan added anomaly, 


dered only slightly UOUL f | tha Ca VEE Yy tui 
except for a few color graphics, the book i is printe di in an apropos light sepia tone. This is haan a 
coffee cookbook which the author makes no bones about. It does, eee begin with several one o 

He skims through such topics as cultivation, different variet- 


two page “chapters” on various to 
ies, processing, opus - movement oa the globe, roasting, and even the origins of (gasp!) in- 
drepat ation of the different styles of coffee drinks 


tor 


stant coffee. From tt 
(cappuccino, Turkish, café latte etc.) ia the use of coffee in other drinks. Finally, there are several 


chapters on food prepared with coffee including: cakes, ice cream, pastries, and other desserts. All is 
mments on the role of coffee in history, politics, culture, and social custom. 


interspersed with ¢ 
» way of example here is one of my favorite quotes from the book: “The small coffee ae met 
at 


its greatest success during the industrialization when beer, which until then had b 
work and even at breakfast, was gradually being pushed aside by coffee. This brought forth a more 
productive labor force, which was better protected when handling machinery and worked more ef - 


ficiently.” Regardless of depth of knowledge anyone interested in the bean will enjoy this delightful 
Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. 


bli Robert]. George, Botanical 


publication, 


reeorge@britorg. 


SIDA 22(1): 144. 2006 


AMELICHLOA: A NEW GENUS IN THE STIPEAE (POACEAE) 


Mirta O. Arriaga Mary E. Barkworth 
Herbario Intermorntain Herharilsm 
Museo Argentino de ie: (as aaIIGIES Department of Biology 
ernar dino R Utah State University 
C1405DJR Buenos Aires, ARGENTINA Logan, Utah 84322-5305, U.S.A. 
ABSTRACT 
A new genus of Sti} P Amelichloa, is described. It differs f ther genera in the tribe in 


the woody, sharp tips of it basal leaves, the presence of smooth longitudinal ribs on its caryopses, its 
persistent stylar bases gnc in the edpent pie of cleistogamous panicles in the axils of its basal 
leaf sheaths. TI ambigua, A. brachychaeta, A. brevipes, A. caudata, 


and A. clandestina. The fie four sneciee are native to South America; A. clandestina is native to 
northern Mexico. A key to the ee is ia ided. Three of them, A. brachychaeta, A. caudata, and A. 
clandestina, have becom ished in disturbed areas on other continents. The name honors Dr. 


Maria Amelia Torres. 


RESUMEN 


Se do Ameliehloe: un nuevo genero pan la ube onpeae (Poaceae). El cu ono difiere a otros 


presentar a muy 
longitudinales e en la supe del cariopse y fee ere persistentes, asi como en la frecuente 
presencia de nos en las axilas de las hojas basales. El género se encuentra constituido por 


cinco Spo A. ambigua, A. Oe ae on Prey e! A. Eauace y A. sain esas Las cuatro 


r . te 
Tres de | ies A. brachychaeta, A.caudata y A.clandesti | blecid A li bad 


| i Se dedica este géneroa la Dra. Maria Amelia Torres. 


This paper arises from the need to determine the appropriate generic treatment 
of three grass species introduced in the United States. In traditional generic treat- 
ments they are known as Stipa brachychaeta, S. caudata, and S. clandestina. All 
three have names in Achnatherum (Barkworth 1993; Jacobs & Everett 1996). The 
first two have, in addition, names in Jarava (Penailillo 2002). Soreng et al. (2003) 
treated S. brachychaeta and S. caudata as species of Jarava and S. clandestina as 
a species of Achnatherum. 

We reviewed fresh material, herbarium samples and type specimens of all 
three species and became convinced that, not only do they all belong to the 
same genus, but that they do not belong to any currently recognized genus. In 
this paper, we present the new genus, explain how it differs from other Ameri- 
can genera, list the species that it includes, and provide a key to the species. 

Etymology.—We are naming the new genus Amelichloa in honor of Dr. Maria 
Amelia Torres who has generously shared with us her extensive knowledge of 
South American Stipede. 


SIDA 22(1): 145 — 149. 2006 


146 BRIT.ORG/SIDA 22(1) 


Amelichloa Arriaga & Barkworth, gen. nov. Type: Amelichloa ambigua (Speg,) Arriaga & 
Barkworth (Stipa ambigua Speg.). 


Species Amelichl liff | li Stipearum laminis apicibus rigidisimis 
a ae 


| i pag dorsali et duobus costis 
1:} ] : 1; 1G] . : Pec SPs eee We fal; 
oO 


et acutissimis 
) 121 * 


basalium 


quam inter species alias in triby pro ai te one 
Plants perennial, cespitose. Culms erect, with 2-3 nodes. Leaves mostly basal; 
sheaths open, smooth, glabrous; auricles absent; ligules scarious, rounded to 
acute; blades stiff, involute, apices very stiff, sharply pointed. Inflorescences 
paniculate, main panicle terminal, apparently wholly chasmogamous, reduced 
cleistogamous panicles often present axillary to the basal leaf sheaths. Spike- 
lets of terminal panicles with | floret, disarticulating above the glumes and be- 
low the floret. Glumes exceeding the floret, acute to acuminate, 1-5-veined; 
anthoecia fusiform, terete; calluses antrorsely strigose, blunt; lemmas pubescent, 
often more densely and/or more persistently so over the midvein and lateral 
veins, hairs on the lower portion about 0.7-2 mm long, hairs on the distal por- 
tion often longer; crown not developed; awns 1-2-geniculate, scabrous, persis- 
tent; lodicules 3; stamens 3, anthers sometimes all of equal size and more than 2 
mm long, sometimes 1 more than 2 mm long and 2 much shorter, sometimes all 
short; ovaries glabrous, style with 2 branches, united at the base, stigmas plu- 
mose. Caryopses fusiform, with 3 smooth, longitudinal ribs at maturity, stylar 
bases 1-2 mm long, persistent, sometimes eccentric; hila linear, about as long as 
the caryopses. Spikelets of axillary panicles 0.5-1 mm long, glumes thin, shorter 
than the florets, florets unawned or with reduced awns. x = 1]? 

Amelichloa includes five species, four of which are South American. The 
fifth species, A. clandestina, grows in northern Mexico. Some species have also 
become established in other parts of the world. The short anthers are probably 
associated with cleistogamy. A similar variation in anther length is also found 
in the primarily South American genus Nassella. In N. leucotricha (Trin. & Rupr.) 
R.W. Pohl, the proportion of florets with the different combinations varies in 
response to environmental conditions. 

Cattle avoid the species of Amelichloa because of the pointed leaves. The 
frequent presence of cleistogenes also favors establishment of the species in ar- 
eas that are grazed or mown. The combination of the sharp leaves and 
cleistogenes means that members of the genus could become serious problems 
in rangelands. 

The only chromosome count that has been made for the genus is of 2n = 44 
for Amelichloa brachychaeta (Bowden and Senn 1962). None of the species has 
been included in any molecular studies. We shall be initiating a study that in- 
cludes some species of Amelichloa, plus many other South American taxa, in 
summer, 20006. 


ADDIACA ANN RADVINODTU AMECLICUINA 147 


1. Hairs at the top of the lemmas 4-5 mm long, numerous A.ambigua 
1. Hairs at the top of the lemmas up to 2.5 mm, long, sparse. 

2. Awns 27-30 mm; florets 7-9.8 mm A. brevipes 

2. Awns 10-25 mm; florets 4-8 mm. 

3, Mature caryopses with inclined, eccentric stylar bases; lemmas glabrous be- 

tween the midvein and the lateral vein, even at the base, usually also gla- 

brous between the lateral vein a the marginal vein A. caudata 
. Mature caryopses with erect, usually centric stylar bases; lemmas pubescent 

between the lateral and marginal veins on the lower 1/2, usually also be- 

tween the midvein and lateral veins. 

4. Florets 4—-5.5 mm long; awns usually 1-geniculate A. brachychaeta 

4, Florets 5.5-8 mm long; awns usually 2-geniculate A. clandestina 


(eS) 


Amelichloa ambigua (Speg.) Arriaga & Barkworth, comb. nov. Basionym: Stipa 
ambigua Speg,, April 1925. Revista Argent. Bot. 1:27. 
Jarava ambigua (Speg.) Penailillo, Gayana, Bot. 59:30. 2002. TypE: ARGENTINA: PROV. BUENOS 
AIRES: Pdo. Gral. Pinto, “in pratis circa Estacion Iriate,” 24 Nov 1905, Spegazzini s.n. (HOLO- 
YPE: LP ex LPS-2412!; isotype: BAA-2946 (col. Typus!)). 
Pee dusenii Hitchc., Contr. U.S. Natl. Herb. 24:271.1925. Type: ARGENTINA: BUENOS AIRES: plains 
Sierra de la Ventana., 29 Nov 1904, P Duséns.n. (HOLOTYPE: US-1161165; PARATYPE: BAA-3038 
ex Herb. Parodi 5054 (col. Typus!)). 


Distribution.—Argentina and Uruguay (Rosengurtt et al. 1970; Zuloaga et al. 
1994; Torres 1993, 1997), introduced to France (Verloove 2005). 


Amelichloa brachychaeta (Godr.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa 
brachychaeta Godr., Mém. Sect. Sci. Acad. Sci. Montpellier 1:450. 1853. 


Nassella brachychaeta (Godr.) Barkworth, Taxon 39:609. 990) Achnatherum brachychaetum 
r.) Barkworth, Phytologia 74:6. apes Jarava brac! (Godr.) Penailillo, Gayana, Bot. 
59: 5 2002. TyPE: FRANCE: du Por l ‘au Jardin Botanique, Patria ignota, “plantes 
etrangerse-an, Port Juvenal. Anon. igh 22 May pla inane PE (probable): MPU; OM: BAA- 
2979, hojas a & b! (col. typus, fragm. d tipo). 
ce eminens fo. viridis Kuntze, Revis. =e PL 3:37). 888 TYPE: ARGENTINA. SANTA FE: Ceres, 
92, Kuntze s.n. (HOLOTYPE: NY; ISOTYPE: COR 
- vanes Griseb., Symb. Fl. Argent. 298- 290, a TYE: ARGENTINA: Cordoba, PG. 
Lorentz 46,187? (HOLOTYPE: GOET, ISOTYPE: CORD, US 866090! (fragm. ex GOET)). 
Distribution.—Argentina, Chile, Pert and Uruguay (Matthei 1965; Rosengurtt 
et al. 1970; Zuloaga et al. 1994; Torres 1993, 1997). Although Tovar (1993) and 
Brako and Zarucchi (1993) do not mention it, the presence of this species in 
Perti is cited in Caro and Sanchez (1971), Torres (1997), and Verloove (2005). In- 
troduced to the U.S.A., Australia Jacobs & Everett 1993), France, and Spain 
(Verloove 2005). 
Amelichloa brevipes (E. Desv.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa 
brevipes E. Desv, Fl. Chil. 6:282. 1854. 
as brevipes (E.Desv.) Penailillo, Gayana, Bot. 59:30. 2002. Type: CHILE: en las provincias 
ntrales de la Republica, C. Gay 1106. s.d. (HOLOTYPE: P; ISOTYPE: BAA! (col. typus fragm 
ues (fragm.)). 


148 BRIT.ORG/SIDA 22(1) 


Stipa hirtiflora Hack. in Dusén, Ark. Bot. 7:5, t.3, (f.7), 7, CL. 4-5). 1908. TYPE: ARGENTINA. SANTA 
Cruz: Arroyo Pelque, Jan 1905, P. Dusén 5640 (HOLOTYPE: W, ISOTYPE: US 1161168, 3168627 
(fragm.)). 

Distribution.—Argentina (Roig in Correa 1978; Torres 1993; Zuloaga et al. 1994), 

Chile (Matthei 1965). 


Amelichloa caudata (Trin.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa caudata 
Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1:75. 1830 


A] th 


tum (Trin.) S.\W.L. Jacobs & J. Everett, Telopea 6:582. 1996. Jarava caudata (Trin.) 
Penailillo, Gayana, Bot. 59:30-31. 2002. TYPE: Specimena Chilensia, J. Lindley s.n. (HOLOTYPE: 
LE-TRIN 1392.01 (& fig); IsoTYPE: BAA 3001! (col. typus fragm), US 2489479 (fragm)). 

Stipa bertrandii Phil. Linnaea 33(3-4):283. 1864. Type: CHILE: Andes de Santiago, Bertrand s.n 
(HOLOTYPE: SGO-PHIL-102; IsoTYPE: BAA 2972! (col. typus frag. ex holotype), SGO 62787, 
63156; US 866140). 

Stipa amphicarpa Phil. Anales Mus Nac. Santiago de Chile 11, lam.3, f.2. 1892. Type: CHILE: prope 
Angol, Dec 1887, R.A. Philippi (HOLOTYPE: SGO-PHIL 99, BAA 2952! (col.typus fragmento ex 
SGO); ISOTYPE: SGO 37393, 62793, US A866145). 

Stipa litoralis Phil. Anales Univ. Chile 93:717. 1896. Type: CHILE: Isla Quiriquina, Mar 1880, R.A 

Philippi s.n. (HOLOTYPE: SGO 62798; US 825120 (fragm. ex SGO)) 


Distribution.—Argentina, Chile and Uruguay (Matthei 1965; Rosengurtt et al. 
1970; Zuloaga et al. 1994; Torres 1993, 1997); introduced to Australia Jacobs & 
Everett 1993), England, Italy, Spain (Verloove 2005), and U.S.A. 


Amelichloa clandestina (Hack.) Arriaga & ees comb. nov. BASIONYM: Stipa 
clandestina Hack., Repert. Spec. Nov. Regni Veg. 8:516. 1 


Achnath ‘land (Hack.) Bark worth, Phytologia 74:6. 1993. TYPE: MEXICO: COAHUILA: 
saltillo, Mar 1908, Arséne aan ECTOTYPE: W, ISOLECTOTYPE: MO 845904t US 1030718, 3168626 
(ragm. ex W). Lectotype designated by Hitchcock, Contr. U.S. Natl. Herb. 24:238. 192 


Distribution.—México; introduced to U.S.A. Hitchcock (1925) included a 
pittieri in S. clandestina and, as a result, included Colombia in the range of S. 
clandestina. The type of S. pittieri (US 531634!), however, belongs in Nassella, as 
is reflected in the combination N. pittieri (Hitche.) Penailillo (1998). Thus, it 
appears that A. clandestina is restricted to northern México. 


ACKNOWLEDGMENTS 


We thank Maria Amelia Torres for her ongoing interest in the tribe, her will- 
ingness to share her knowledge, and her well documented publications. We also 
thank the curators of BAA, CDA, LP, MO, and US for making their specimens 
available to us for this and our other studies in the Stipeae, J. Landers and J. 
Valdés Reyna for bringing the presence of A. clandestina to Barkworth’s atten- 
tion, RJ. Soreng for clarifying Hitchcock's attribution of Stipa clandestina to 
Colombia, and G. Davidse and S.WL. Jacobs for their careful review and thought- 
ful comments. 


ARDIAGA AND RARPKUIWOARTH AMELICHINA 149 


REFERENCES 


BarkworthH, M. 1993. North American Stipeae (Graminae): taxonomic changes and other 
comments. Phytologia 74:1-25. 

Bowoen, W.M. and H.S. Senn. 1962. Chromosome numbers in 28 grass genera from South 
America. Canad. J. Bot.40:1115-1124. 

Brako, L.and J.L. ZaruccHi. 1993. Catalogue of the flowering plants and Gymnosperms of 
Peru. Monogr. Syst. Bot., Missouri Bot. Gard. 45. 

Brown, W.V. 1952.The relation of soil moisture to cleistogamy in Stipa leucotricha. Bot. Gaz. 
113:438-444. 

HitcHcock. A.S. 1925. North American species of Stipa. Contr. U.S. Natl. Herb. 24:215-262. 

Jacoss, S.W.L.and J. Everett. 1996. Austrostipa,a new genus, and new names for Australasian 
species formerly included in Stipa (Gramineae). Telopea 6:579-595. 

MattHel, O.R. 1965. Estudio critico de las gramineas del género Stipa en Chile. Gayana, Bot. 
13:1-137. 

PENAILILLO, P. 1998. Nuevas combinacidnes en el género Nassella E. Desv.emend.Barkworth 
(1990) (Poaceae, Stipeae). Gayana, Bot. 55:85-88. 

PENAILILLO, P. 2002. El género Jarava Ruiz et Pavon (Stipeae-Poaeae): delimitacion y nuevas 
combinaciones. Gayana, Bot. 59:30. 

Roic, FA. 1978. Género Stipa L. In: Correa, Flora Patagénica, Coleccidn Cientifica del INTA, 
7(3):288-327. 

Rosencurrt, B., B. ARRILLAGA DE Marrel, and P. Izacuirre De Artucio. 1970. Gramineas Uruguayas. 
Univers. de la Republica, Depto. Publicacidnes. Montevideo, Uruguay. 

SoRENG, R.J., BM. Peterson, G. Davipse, E.J. Jupziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 
2003. Catalogue of New World grasses (Poaceae): IV. Subfamily Pocideae. Contr. U.S. 
Natl. Herb. 48:1-730. 

Torres, M.A. 1993.Revision del género Stipa (Poaceae) en la Prov.de Buenos Aires. Comision 
de Investigaciones Cientificas, Prov. Buenos Aires, Monografia 12:1-62. 

Torres, M.A. 1997. Stipa (Gramineae) del noroeste de la Argentina. Comisién de 
Investigaciones Cientificas, Prov. Buenos Aires, Monografia 13:42-65. 

Tovar, O. 1993.Las Gramineas (Poaceae) del Peru. Ruizia 13:1-480. 

Vertoove, F. 2005.A synopsis of Jarava Ruiz & Pav.and Nassella E. Desv. (Stipa L.,s./.) (Poaceae: 
Stipeae) in southwestern Europe. Candollea 60:97-117. 

ULOAGA, F.O., E.G. Nicora, Z.E. RUGOLO DE AGRASAR, O. Morrone, J. PENSIERO, aNd A.M. CIALDELLA. 1994. 
Catalogo de la Familia Poaceae en la Republica Argentina. Monogr. Syst. Bot., Missouri 
Bot. Gard. 47:1-178. 


150 BRIT.ORG/SIDA 22(1) 


Book NOTICES 
Flora of North America (Asteraceae) 

Fiora oF Nort AMERICA EpitoriAL Committee (Eds.) 2006. Flora of North America 
North of Mexico. Vol. 19. Magnoliophyta: Asteridae, part 6: Asteraceae, 
part 1. (ISBN 0-19-530563-9: 978-0-19-530563-0, hbk.). Oxford Univ. Press, 
198 Madison Avenue, New York, NY 10016, U.S.A. (Orders: http:// 
www.oup.com). $95.00, 579 pp., illustrations, maps, 8 1/2" x 11". 

FLora or NortH AMeriIca EpttortAL Committee (Eds.) 2006. Flora of North America 
North of Mexico. Vol. 20. Magnoliophyta: Asteridae, part 7: Asteraceae, 
part 2. ISBN 0-19-530564-7; 978-0-19-530564-7, hbk.). Oxford Univ. Press, 
198 Madison Avenue, New York, NY 1OO16, U.S.A. (Orders: http:// 
www.oup.com). $95.00, 666 pp., illustrations, maps, 8 1/2" x 11" 

Frora oF NortH America Epiroriat Committee (Eds.) 2006. Flora of North America 
North of Mexico. Vol. 21. Magnoliophyta: Asteridae, part 8: Asteraceae, 
part 3. ISBN 0-19-530564-4; 978-0-19-530565-4, hbk.). Oxford Univ. Press, 
198 Madison Avenue, New York, NY 10016, U.S.A. (Orders: http:// 
www.oup.com). $95.00, 616 pp., illustrations, maps, 81/2" x 11" 


From FNA website.—Covering the Asteraceae, or Composite, family, the largest family in the Flora of 
North America area, these three volumes contain ne cae 420 species classified in +18 genera and 14 
tribes. More than 60% of the s > the flora region and almost 10% are known as 
introductions that have miatanaticedl Members of ‘his tamily of higher plants can be found to flour- 

ish in many habitats. Sunflowers, daisies, goldenrods, sagebrush, asters, zinnias, arnicas, chamomile, 
cory, lettuce, coneflowers, ragweeds, tickseeds, thistles, dandelions, 


> 


chrysanthemums, marigolds, ch 
tarweeds, cockle dee ereen and many other familiar plants are members of the Asteraceae. 
Each of the genera within these volumes has one or more representative species illustrated with a 
line drawing; more than 660 species are illustrated. Occurrence maps for nearly 3,300 species anc 
accompany the identification keys, descriptions, and discussions about these 


subspecies, b 


plants. 


SIDA 22(1): 150. 2006 


TAXONOMY OF THE CUSCUTA PENTAGONA COMPLEX 
(CONVOLVULACEAE) IN NORTH AMERICA 


Mihai Costea (corresponding author Guy L.Nesom 
Department of Biology Botanical Research Institute of Texas 
fa a pallies! University can Stre 
5 | West Fort Worth, Texas 76102- He U.S.A. 
Waterloo, Ontario esom@brit.org 
N2L 3C5, CANADA 


mceostea@wlu.ca 
Sasa Stefanovic 
Department of Biology 
University of Toronto at : ississauga 
359 Mississauga Road 
Mississauga, cakes 
L5L 106, CANADA 


ABSTRACT 


Cuscutd pentagona, C. campestris, and C. glabrior are closely similar taxa recognized here at specific 
a 8 y P 
rank. Closely related to these are C. runyonii, C. harperi, C. plattensis, and C. sandwichiana, as well as 


G: pu UoNG var. cle an and C. polygonorum. Cuscuta runyonii, C. harperi, ane Ss ea aan 


L 
are relatively narrow € : pla ttensisand C.¢labrior have wider but ay parently 


while C. obtusiflora var. ¢ Pagid C. polygonorum, and especially C. pentagona ae Ee Peierls 
are widespread and br oadly sympatric in North America, at least in some regions. Cuscuta campestris 
is one of the most widespread species in the world, being additionally recorded from South America, 


Europe, Asia, Africa, and eee Comparisons among these nine species are presented as a key 


and as comparative d M images are provided for flowers, fruits, seeds, and pollen. 


RESUMEN 


Cuscutd ponapane: . canon y GC. ais son taxa fuertemente semejantes, reconocidos aqui 
como especies as estan C. runyonii, C. harperi, y C. Ss 
como tambien G; gps ite var, eagles yiG: pono Cuscuta runyonii, C. ee 


C. plattensis y C.glabrior tienen ran 


I 
14 di 


8 
eee Cn ca tusiflora var.glandu losa, a 


y especialmente G pemieepng y.G. eee son comunes y eae iamente Sue en 
d 


Norteameéric siendo 


mencionada a Sudamérica, Europa, Asia, Arica y Australia Las comparaciones entre estos seis 


Se ofrecen imagenes de SEM de las 


I E 


f 


ores Hates semillas y polen. 


Engelmann (1859) distinguished three varieties of Cuscuta pentagona Engelm. 
in addition to the typical form: var. calycina, var. verrucosa, and var. microcalyx. 
Yuncker (1932, 1965) treated var. calycina and var. verrucosa at specific rank 
but provided a new name for each: C. campestris Yuncker (nom. nov.) and C. 


SIDA 22(1 


= 


> 151-175. 2006 


152 BRIT.ORG/SIDA 22(1) 


glabrior (Engelm.) Yuncker (comb. nov.), respectively. He treated var. microcalyx 
as a synonym of typical C. pentagona. Recent studies have not recognized C. 
campestrisat any rank (e.g, Beliz 1986; Gandhi et al. 1987), while C. glabrior has 
been regarded asa variety of C. pentagona (Gandhi et al. 1987) or as nomencla- 
tural synonym of C. indecora Choisy var. neuropetala (Engelm.) Hitche. (Beliz 
1986). Three other species from North America, C. harperi Small, C. plattensis 
A.Nels.,and C. runyonii Yuncker, were included by Yuncker in the same group 
(subsect. Arvenses Yuncker) with C. pentagona and C. campestris (Yuncker 1965). 
Although C. runyonii has been considered a good species by all authors, C. 
harperi and C. plattensis were treated by Beliz in her Ph.D. dissertation (1986) 
as nomenclatural synonyms of “C. pentagona var. polygonorum’ and C. indecora, 
respectively, 

The current paper is the second of several to present taxonomic analyses 
of Cuscutad species groups and complexes. The first (Costea et al. 2005) gave an 
assessment of Cuscuta nevadensis and Cuscuta veatchii; three others follow in 
this issue (Costea et al 2006a,b,c). Other two papers that are now in preparation 
will deal with the phylogeny of Cuscuta and its infrageneric taxonomy, includ- 
ing the formal delimitation of the groups of species treated in the four papers 
published in this issue. Evaluations of the status of all these taxa are necessary 
for the accounts of Cuscuta in the Flora of North America and the forthcoming 
revision of The Jepson Manual (California). The comments below on reproduc- 
tive biology and taxonomy are in perspective of the whole genus. 


pes) 


— 


REPRODUCTIVE BIOLOGY AND TAXONOMY OF CUSCUTA 
Scarcity of information regarding the reproductive biology of Cuscuta species 
limits progress toward understanding the taxonomy and evolution of this ge- 
nus. For example, a detailed study of floral biology is available only for C. 
attenuata Waterfall (Prather & Tyrl 1993), which is very closely related to to C. 
indecora. Although they may be sympatric over an extensive geographical 
range, species such as C. campestris and C. pentagona have never been found 
“growing together at the same site” (Musselman 1986). Apparently a single her- 
baceous host cannot support more than one parasite (often a single Cuscuta 
plant simultaneously parasitizes several different host plants), although it does 
not seem impossible that colonies of different Cuscuta species might exist side 
by side. Furthermore, although some species (e.g., C. epithymum Murray, C. 
rostrata Shuttlw. ex Engelm. &@ A.Gray) have been reported to be pollinated by 
insects (Beliz 1986; Musselman 1986; Holm et al. 1997), others such as C. 
pentagona and its relatives apparently are autogamous (Verdcourt 1948; Beliz 
1986; Musselman 1986; Prather & Tyrl 1993; Dawson et al. 1994). In the C. 
pentagona complex, we observed that anthers dehisce longitudinally before 
{lowers open, further substantiating previous observations of autogamy. The 
low pollen/ovule ratio (= 65) found by Beliz 1986) in C. pentagona Gncluding C. 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 153 


campestris) supports the idea that autogamy may be common (see Cruden 1977). 

The occurrence of inbreeding may help explain the relatively high num- 
ber of taxa (species and varieties) that are recognizable as morphological enti- 
ties in Cuscuta. The existence of intermediate-like plants observed rarely in 
populations of the C. pentagona complex probably reflects populational vari- 
ability in closely related taxa. Hybridization also is a possible explanation, but 
experimental attempts to produce hybrids (e.g. between C. indecora and C. 
attenuata) have failed (Prather & Tyrl 1993). 

Seed dispersal of many Cuscuta species seems unspecialized (Kuijt 1969). 
“Natural” means of seed dispersal are unknown. Lyshede (1984) suggested that 
wind may play a significant role in dispersal because of the pits present on the 
seed coat of dry seeds. However, Cuscuta seeds do not possess “classical” adap- 
tations for wind dispersal, and the alveolate/papillate seed coat seems more an 
adaptation related to the germination process (Costea & Tardif 2006). Kuijt 
(1969) mentioned that seeds remain viable while they pass through the diges- 
tive system of sheep, but Gdmez (1994) in Sierra Nevada, Spain, reported that 
hosts infested with C. epithymum were avoided by the Spanish ibex, the main 
herbivore in the area. Herbivores may avoid plants infested with Cuscuta be- 
cause of the yellowish color (Costea & Tardif 2006). Capsules and seeds of 
Cuscuta float, and although water dispersal has never clearly been documented, 
it may occur only in the species that prefer the vicinity of water, such as C. 
gronovii Willd.ex J. A. Schult. Dispersal over long distances of weed species such 
as C. campestris has been connected with anthropomorphic activities. The prin- 
cipal means of world-wide dispersal of Cuscuta weeds has been through con- 
taminated seeds of forage legumes (alfalfa, clover, and lespedeza, especially Les- 
pedeza cuneata (Dumont) G. Don; reviewed by Dawson et al. 1994; Costea & 
Tardif 2006). Furthermore, when wetted, the seed coat in Cuscuta species be- 
comes papillose-gelatinous and adheres easily to soil particles, and seeds may 
thus be carried by farm machinery (Costea & Tardif 2006). 


METHODS 


Descriptions of morphology and micromorphology are based mostly on speci- 
mens from herbarium NY, which includes Yuncker’s herbarium (Appendix 1). 
Measurements of floral parts except anthers were done on rehydrated her- 
barium material. Length of flowers was measured from the base of calyx to the 
tip of straightened corolla lobes. Pollen, anthers, seeds, and capsules were mea- 
sured on SEM pictures taken with a scanning electron microscope Hitachi S- 
570 at 15 KV. Samples were coated with 30 nm gold using an Emitech K 550 
sputter coater. Standard terminology for cell types and surface sculpturing pat- 
terns follows Barthlott et al. 1998); pollen terminology follows Hoen (1999). 
Conservation status was assessed using NatureServe (2005) ranks and criteria. 

Identification of Cuscuta often isa lengthy process because rehydration of 


154 BRIT.ORG/SIDA 22(1) 


flowers, dissection, and examination under a microscope are usually necessary. 
Variation may be expected on the same individual (e.g, Gandhi @ Thomas 1983; 
Severova 1991). Furthermore, because flowers do not mature simultaneously, 
the size and ratios between different floral parts on the same plant may vary to 
some extent. A range of flowers, fruits, and seeds belonging to the same plant 
should be examined. The angled calyx of C. pentagona and C. harperi is best 
ower and during early fructification; full development of capsules 


— 
—_— 


observed inf 
may obscure the character. 


MICROMORPHOLOGY 


Flowers and capsules.—Laticifers, papillae, and multicellular protuberances may 
occur on the floral parts and capsules and distinctions between these features 
can be important in recognizing members of the C. pentagona complex (mag- 
nification of LOOX or more recommended). Laticifers are visible in the calyx 
and sometimes in the corolla and capsules of all taxa. Yuncker usually referred 
to them as “pellucid glandular-appearing cells” (Yuncker 1921, 1932, 1965). La- 
ticifers may appear isolated, rotund, ovoidal or tangentially elongated, or they 
can be organized in rows (Fig. 1 c,e). Laticifers from the stems are perivascular, 
multinucleate, and articulated (Lyshede 1985). Papillae are more or less radi- 
ally elongated cells, and they are present in the calyces, corollas, and capsules 
of C. harperi, C.glabrior, and C. runyonii (Fig. Le,f; 2. a,b,c). Multicellular protu- 
berances are larger and they may occur in C. runyonii along the midveins of the 
calyx lobes (Fig. 2c). Epicuticular wax on the perianth, when present, is repre- 
sented by longitudinally reticulated rodlets (Fig. Lf). 

Pollen.—Pollen is relatively uniform among species. Grains are 3(-4)- 
zonocolpate (rarely +-6 loxocolporate in aberrant grains),and they may be poly- 
morphic in the same anther or flower, from spheroidal to prolate (Fig. 3). Orna- 
mentation varies from tectum imperforatum or witha few puncta in C. glabrior, 
C.runyonii,and C. plattensis, to tectum perforatum in C. pentagona, C. campestris, 
C. harperi, and C. obtusiflora var. glandulosa (Fig. 3). The latter three species 
have sexine often persistent in punctum apertures (Fig. 3). Cuscuta pentagona 
and C. obtusiflora var. glandulosa have the largest puncta, up to 0.7 fm in diam- 


—_— 


= 


eter, sometimes approaching a microreticulate ornamentation (Fig. 3). 
Seeds.—A striking feature of the seed coat epidermis is that when seeds 
dry, cells of the outer cell wall invaginate, which causes the seed surface to be- 
come alveolate (Fig. 4; see also Lyshede 1984; Knepper et al. 1990; Costea & Tardif 
2006). Water uptake induces bulging of the invaginated epidermal walls, and 
epidermis cells become swollen and papillose (Fig. 4c). The diameter of seed 
coat epidermal cells is 17-50 pm in C. pentagona, C. campestris, C. glabrior, C. 
runyonii, and C. harperi and 15-20 um in C. plattensis, C. obtusiflora var. 
glandulosa, and C. polygonorum. The seeds of C. runyonii and C. polygonorum 
have epicuticular wax organized as longitudinal rodlets (Fig. 4d), a feature not 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 


Fic. 1.a. Calyx of C. campestris (scale bar = 1 mm);b. Calyx of C. p (scale t 0.5 
ey | 1 


capsule of C. ); d. Flower of C. (scale | 0.75 mm); e. Flower of C. 


bar = 0.5 mm); f. P pill th lla lob fC harperi (scale bar = 0.43 um). 


with 


ithma 
arperi (scale 


156 BRIT.ORG/SIDA 22(1 


= 


Fic. 2. a. Flower with maturing capsule of C. gigenes (scale ae 1 mm); b. panies on the capsule of C. glabrior (scale 
bar = 0.75 Lim); c. Flower of C. runyonii projections; d. Flower of C. plattenis (scale bars = 
1mm). 


observed in the other species. The hilum is round to broadly elliptic; the cells 
are smaller than in the rest of the seed coat and they are radially oriented around 
the vascular scar of the funiculum (Fig. 4 b,d). The vascular scar is linear, ob- 
lique to vertical (on broadest diameter of the seeds), 0.025-0.09 mm in C. 
pentagona, C.campestris, C.glabrior, C. pe C. harperi,and C.sandwichiana 
and 0.125-0.25 mm in C. plattensis, C. obtusiflora var. glandulosa, and C. 
polygonorum (Fig. 4 ae,f). 


TAXONOMY 


This g er oup of species is characterized by globose or mostly depressed-glo 
capsules with relatively large interstylar apertures, which sometimes are thick- 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 157 


eee rectum Sanu but sexine 


he S Saat of pollen a-b. bua bast grains oe nee «Cp is (scale bar = 8.6 um); d. C. 


BRIT.ORG/SIDA 22(1) 


Fig. 4, Malahat f Is. a. C. campestris, ventral view (scale ba 
ranvoti teal bar = 120 j1m); e, See 
(scale bar = 0.5 mm). 


= 0.5 mm); b. Hilum and alveolate seed coat of C. 
= 150 um); ¢. Ppl on yaa jaa co tof gna Gale a = a yale d. Hilum of ¢. 
dof © harp 


C. plattensis, ventral 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 159 


ened but not raised to form a stylopodium. The corolla is persistent at the base 
of the capsules or surrounding them. 

Cuscuta sand wichiana, a Hawaiian endemic, was placed by Yuncker (1932) 
in subsect. Californicae because it exhibits a reduction of the infrastaminal 
scales similar to the other species (C. sandwichiana was omitted in his 1965 treat- 
ment). This species, however, does not appear to be evolutionarily allied to C. 
californica complex. Beliz 1986) hypothesized that C. sandwichiana has evolved 
from C. cephalanthi Engelm., which is a member of the C. gronovii Willd. ex 
Roem. & Schult. complex. However, the stems of C. sandwichiana grow and 
branch as in species of the C. pentagona complex: the main and secondary stems 
grow continuously and never twine around the host. Instead, tendril-like 
branches produced from axillary buds of the rudimentary leaf-scales fix the 
parasite to the host. In C.gronovii and its relatives (e.g.,C. cephalanthi, C. umbrosa 
Beyr. ex Hook., and C. rostrata Shuttlw. ex Engelm. & A. Gray), haustoria are 
formed as the main stem twines around the host, and no tendril-like axillary 
branches are generated (as first observed by Dawson 1984 in C. gronovii). The 
calyx and corolla of C. sandwichiana are relatively similar in morphology to 
those of C. campestris, although flowers are larger in C. sandwichiana. Seeds of 
C.sandwichiana are commonly angled, like those of C. campestris, and not dor- 
soventrally compressed as in C. cephalanthi. 

Contrary tosome relatively recent treatments (e.g. Austin 1986; Musselman 
1986), C. campestris has not been accepted as a distinct taxon in modern North 
American overviews of the genus (Beliz 1986; Gandhi et al. 1987; Kartesz et al. 
1999; USDA NRCS 2004). This judgment has been based on the allegedly vari- 
able morphology of the calyx, which was the main character used by Yuncker 
to differentiate this taxon from C. pentagona s. str. Beliz (1986) wrote: “The angled 
calyx (of C. pentagonas. str.) is not a constant feature, it is present in some flow- 
ers and absent on others in an individual, ... this feature may be an artifact cre- 
ated when flowers dry out.” In contrast, we have found the angled calyx of C. 
pentagona to bea remarkably constant feature (Fig. 1b). Because the calyx lobes 
of C. campestris are overlapping, the calyx sometimes may appear angled in 
pressed flowers, but if the flowers are hydrated, the calyx usually appears 
“rounded” (Fig. la). The prominent angles of the calyx in C. pentagona are the 
result of a different morphology of the calyx lobes (Fig.1b). Furthermore, small 
but consistent differences in the size of floral parts, pollen, capsules, and seeds 
(see key below) usually separate the two taxa. Size overlap between the two taxa 
does occur, as well as intermediate-like plants, although the latter are uncom- 
mon. Austin (1979) discussed methods of pressing to avoid obscuring calyx traits 
in all species, but particularly in C. pentagona and C. campestris. 

Four years before Yuncker formally recognized Cuscuta campestris as a 
separate species from C. pentagona, he was aware that his taxonomic solution 
might be received with reluctance. Ina letter addressed to WC. Ferguson, accom- 


160 BRIT.ORG/SIDA 22(1) 


panying a collection of C.campestrissent to him for identification (Ferguson 7795, 
NY), Yuncker (1929) wrote: “I suspect I will be accused of various bad tendencies 
in the way of species making, but I think I have clarified the ‘arvensis’ situation 
somewhat.” The assessment presented here also finds that C. campestris is con- 
sistently distinct from C. pentagona, although differences may seem subtle, and 
we believe their recognition as species best reflects the biological situation. 

Similarly, Cuscuta campestris may be sometimes difficult to distinguish 
from C. glabrior, but the two taxa can be usually differentiated using a combi- 
nation of features indicated in the key and descriptions. Cuscuta runyonii is 
most similar to C. glabrior, and C. harperi to C. pentagona (see descriptions; 
Yuncker 1932; 1942). Cuscuta plattensis does not appear to be evolutionarily al- 
lied to C. indecora, in contrast to the suggestion by Beliz (1986). Yuncker (1932, 
p. 140) considered C. plattensis to be “very closely related” to C. campestris. The 
morphology of seeds of C. indecora is different from the species of C. pentagona 
complex and more similar to that of C. gronovii and its relatives. 

Two North American taxa previously treated within Cuscuta subsect. 
Platycarpae (Yuncker 1932), C. obtusiflora var. glandulosa Engelm. and C. 
polygonorum Engelm. are included here in subsect. Arvenses on the basis of their 
overall morphological similarity with the other species of this subsection. This 
close relationship was also emphasized by Beliz (1986), who referred to C. 
polygonorum as “C. pentagona var. polygonorum,” within which C. harperi was 
included asa synonym. In contrast, we found that although C. polygonorumand 
C. obtusiflora var. glandulosa apparently are related to the species of subsect. 
Arvenses, each is clearly delimited morphologically. Cuscuta australis R. Br from 
Australia and Asia, whichis known only from one collection in North America 
Jefferson Co., New York, 1926, Muenscher s.n., NY), probably belongs to this 
group of species as well. This latter species has not been included in this taxo- 
nomic treatment limited in scope to North America. 

In the Cuscuta pentagona c th ively narrow endemics 
(C. runyonii, C. harperi, and C. sandwichiana), two with wider but apparently 
discrete ranges (C. plattensis and C. glabrior), and four species (C. pentagona, C. 
campestris, C. polygonorum, and C. obtusiflora var. glandulosa) widely distrib- 
uted and broadly sympatric, at least in some regions. The exact North Ameri- 
can distributions of C. campestris and C. pentagona, particularly, require addi- 
tional floristic study. For additional synonymy see Yuncker (1932, 1965). 


— 


1 


KEY TO SPECIES OF THE CUSCUTA PENTAGONA COMPLEX 
IN NORTH AMERICA 


1. Infrastaminal scales well-developed, densely fringed or fimbriate. 
2. Corolla lobes obtuse, straight 8.C. obtusiflora var. glandulosa 
2, Corolla lobes acute, inflexed at apices. 
3. Calyx angled, loose around the corolla tube, 0.3-1.1 mm long; lobes broadly- 
ovate rhombic, + auriculate, overlapping and forming prominent angles at 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 161 


sinuses; flowers 0.9-2.5 mm one: corolla tube 0.5-1.2 mm; anthers 0.2-0.35 
mm long; capsules 1.9-2.4 Xx 1 
4. Flowers commonly 5-merous, 1.4—2.1(—2.5) mm long; calyx lobes not cari- 
nate and without rows of prominent laticifers along midveins (Fig. 1 b, d); 
seeds usually 4 per capsule, angled, subrotund to broadly elliptic, longitu- 
dinal groove absent 1.C. pentagona 
4. Flowers commonly 4-merous, 0.9-1.1(-1.5) mm long; calyx lobes carinate 
or with rows of prominent laticifers along midveins (Fig. le); seeds usually 
1 per capsule, almost spherical, with a longitudinal groove on the ventral 
face (Fig. 4e) 2.C. harperi 
. Calyx rounded, closely enclosing the corolla tube, 1-2.1 mm long;lobes ovate- 
triangular to triangular, not auriculate, overlapping or not and not forming 
prominent angles at sinuses; flowers 2.1-4.6(-5) mm long, corolla tube 1.1- 
2.5 mm; anthers 0.4-0.7 mm long; capsules 1.3-2.8 X 1.9-3.8 mm 
5. Calyx lobes about as wide as long, evidently overlapping (Fig.1a); persis- 
tent corolla enveloping 1/3 or less of the capsule (the basal portion) (Fig. 
1 


Ww 


C). 3.C. campestris 
5. Calyx lobes longer than wide, not rei or only slightly so; persis- 
tent corolla enveloping 1/2 or more of the capsule. 
6, Flowers (whole flowers or sometimes ee mae lyx) and 


re papillate; reddish-brown when dried, corolla erpandiic. alehere 
cate between the lines of stamen attachments; infrastaminal scales 
ing corolla tube; seeds an 
7. Calyx without Sane Bee (Fig. 2a) 4.C. glabrior 
7. Base of calyx corresponding to each lobe with a reflexed spur-like 
multicellular projection 0.1-0.6 mm long, and often with smaller mul- 
ticellular projections on the midveins of calyx lobes (Fig. 2c) 
5.C. runyonii 
6. Flowers (calyx, corolla, ovary) and capsules not papillate, creamy white 
when dried; corolla tube narrowly campanulate to cylindric-campanu- 
late, not saccate between the lines of stamen attachments (Fig. 2d); 
infrastaminal scales shorter than corolla tube; seeds dorsoventrally 
compressed (Fig. 4f) 6.C. plattensis 
1. Infrastaminal scales absent or reduced, bifid, or with a few distal teeth or fimbriae. 
8. Flowers (4-)5-merous, 3-4(—5) mm;infrastaminal scales absent or reduced, trun- 
cate or triangular, bifid or distally with a few teeth or fimbriae; endemic to 
Hawaii 7.C, sandwichiana 
8, Flowers (3-)4-merous, 2—2.7 mm; infrastaminal scales oblong, mostly shallowly 
bifid with 1-3 fimbriae on each side of the filament attachment or a few irregu- 
lar fimbriae at the apex; widespread distribution in N America 9.C. polygonorum 


|. Cuscuta pentagona Engelm., Amer. J. Sci. Arts 43:340. 1842. Cuscuta arvensis Beyr. 
?ngelm. var. pentagona (Engelm.) Engelm., Trans. Acad. Sci. St. Louis 1:494. 1859. LECTO- 
TYPE (Yuncker 1932): U.S.A. VIRGINIA: Norfolk, “On Euphorbia or Tragia,” 1849, Rugel s.n. (MO). 


Cuscuta pentagona Engelm. var. microcalyx Engelm., Amer J. Sci. 45:76.1843. LECTOTYPE (Yuncker 1921): 
U.S.A. ILLINOIS. [Cass Co.| Beardstown, 1842, Geyer s.n. (MO). Engelmann cited only “Illinois.” 

Cuscuta arvensis Beyr. ex Engelm. in A. Gray, Man. Bot. (ed. 2) 336. 1856. Epithymum arvense (Beyr. 
ex Engelm.) Niewl. & Lunell, Amer. Midl. Naturalist 4:51. 1916. Type: Engelmann cited only “(in 
herb. Berlin),” perhaps referring to a Beyrich collection. As noted by Yuncker (1921, p.51),a MO 
collection by Beyrich in 1845 from North Carolina may be “the type?” (an isotype). Evidently 


162 BRIT.ORG/SIDA 22(1) 


Engelmann intended C. arvensisasa broad concept variants might be iden- 
tified as C. pentagona, C. pentagona var. microcalyx, C. pentagona var. calycina, or C. verrucosa. 
Stems |.4-2.1(-2.5) mm in diameter, yellow to orange. Inflorescences dense, 
corymbiform to glomerulate of 3-15(-20) flowers; pedicels 0.5-3(-4.5) mm; 
bracts | at the base of clusters and 0-1 at the base of pedicels, 0.4-2.2 x 0.2-13 
mm, membranous, ovate, ovate-triangular to lanceolate, margins entire, apex 
acute. Flowers (4-)5-merous, 1.4-2.1(-2.5) mm (Fig. 1d), membranous, whitish 
when fresh, yellowish to brown when dried; papillae sometimes present on the 
corolla lobes; laticifers evident in the calyx and less obvious in the corolla and 
ovary/capsule, isolated or arranged in rows, ovoid or elongated; calyx yellow to 
brown, * reticulate, shiny or not, angled, cupulate, ca. as long as the corolla 
tube, divided 1/2-2/3 the length, lobes overlapping and forming prominent 
angles at sinuses, broadly-ovate to rhombic, auriculate, not carinate, margins 
entire, apex rounded (Fig. 1b); corolla persistent; tube campanulate, 0.7-1.2 mm, 
not saccate between the lines of stamen attachments; lobes spreading, 0.65-1.1 
mm long, triangular-lanceolate, margins entire, apex acute to acuminate, in- 
flexed; stamens exserted, shorter than corolla lobes, anthers subrotund to 
broadly elliptic, 0.25-0.30 x 0.20-0.28 mm, filaments 0.3-0.4 mm; pollen grains 
18-25.2 um long; tectum perforatum; maximum diameter of puncta 0.6 pm; 
supieicelal Sea eae to conical + uniformly distributed (Fig. 3e); 
rolla tube, oblong-ovate, rounded, + uniformly, 
dense fimbriate; styles everily filif orm, O./-1.1 mm, ca.as long as the ovary; stig- 
mas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose- 
depressed to ovoid, 1.9-2.4 x 1.6-2.5 mm, not thickened or raised around the 
medium to large interstylar aperture, translucent or not, persistent corolla at 
the capsule base. Seeds 4 per capsule, angled, subrotund to broadly elliptic, 0.9- 
Ll x 0.8-1 mm, seed coat cells reticulate to alveolate/papillate, 18-35 tm in 
diameter, epicuticular wax absent; hilum region round 0.4-0.5 mm in diam- 
eter, vascular scar 0.07-0.09 mm long, vertical to slightly oblique. 2n = 56 
(Fogelberg 1938), ca. 44 (Pazy & Plitmann 1995). 

Distribution and ecology.—CANADA: Manitoba (based on only 2 collections 
from DAO and MTMG respectively). U.S.A.: Alabama, Arkansas, D.C., Delaware, 
Florida, Georgia, Kansas, Illinois, Indiana, Maryland, Massachusetts, Michigan, 
Minnesota, Mississippi, Missouri, Montana, New Jersey, New York, North Caro- 
lina, North Dakota, Oklahoma, Pennsylvania, South Carolina, South Dakota, 
Tennessee, Texas, Virginia, Washington. We have seen no evidence that C. 
pentagona has spread outside of North America, where it is less common than 
C.ca mpestris. Isolated European records identified as this taxon (e.g, Buia 1938) 
are instead C. campestris. Flowering Jun-Nov. Hosts: numerous hosts (Gaertner 
1950) growing in a wide variety of climates and soil types. 

Conservation status.—G4G5 (apparently secure to secure) (G5, NatureServe 
2005). 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 163 


2. Cuscuta harperi Small, Fl. S.E. US. ed. 2, 1361, 1375. 1913. Type: US.A. Georcia: 

Altamaha Grit, between Peacocks and Harrison, on Chond rophora virgata, 20 Jul 1906, Harper 

s.n. (HOLOTYPE: NY, see Yuncker 1943; ISOTYPE: NY). 
Stems 0.15-0.3 mm in diameter, orange-yellow. Inflorescences loose, 
corymbiform of 2-10(-15) subsessile or short pedicellate flowers; pedicels 0.5- 
2.5(-3) mm; bracts | at the base of clusters and absent (rarely present) at the 
base of pedicels, like those of C. pentagona. Flowers 4(-5)-merous, 0.9-1.1(-L5) 
mm (Fig. le), fleshy, creamy-white when fresh, yellow to brownish when dried; 
papillae present on the corolla lobes (Fig. Lf); laticifers evident along midveins 
of calyx and less obvious in the corolla and ovary/capsule, isolated or in rows, 
elongated; calyx yellow-brownish, + reticulate, rarely shiny, about as long as 
the corolla tube, angled, cupulate, lobes basally overlapping and forming promi- 
nent angles at sinuses, broadly-ovate rhombic, + auriculate, + carinate or with 
laticifers along midveins, margins entire, apex rounded; corolla persistent; tube 
campanulate, not saccate, 0.5-0.7 mm; lobes erect to reflexed, 0.4-0.7 mm, tri- 
angular-ovate, margins entire, apex subacute to acute, inflexed; stamens ex- 
serted, slightly shorter than corolla lobes, anthers subrotund to broadly ellip- 
tic, 0.2-0.25 x 0.15-0.18 mm, filaments 0.16-0.25 mm; pollen grains 17.5-25 um 
long; tectum perforatum but sexine often persistent in puncta aperture; diam- 
eter of puncta 0.4 um; supratectal process granular granular to conical + uni- 
formly distributed (Fig. 3 a,b); infrastaminal scales as long as the corolla tube, 
narrowly oblong, rounded, shortly and irregularly-dense fringed in the distal 
half; styles uniformly narrow terete to filiform, 0.5-0.9 mm, shorter than the 
ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, 


globose to ovoid, 1.2-2.3 x 1.2-1.6 mm; not thickened around the small to mod- 
erately large interstylar aperture, almost translucent, with persistent corolla 
enveloping the 1/4-1/3 of the capsule bases. Seeds usually 1-2 per capsule, 
subglobose, with a longitudinal groove on the ventral face, 0.92-1.11 x 0.85-L1 
mm, seed coat cells alveolate/papillate, 35-50 um in diameter; epicuticular wax 
absent; hilum region terminal, round, 0.14-0.16 mm in diameter, vascular scar 
0.025-0.03 mm long, vertical to slightly oblique. 2n = ? 

Distribution and ecology.—U.S.A.: Alabama and Georgia. Flowering Sep- 
Nov. Hosts: Bigelowia nuttallii, Croton willdenowii, Helianthus longifolius, Hy- 
pericum gentianoides, Liatris microcephala, and other species of sandstone 
outcrops (see also Yuncker 1943). 

onservation status.—G1G2 (critically imperiled to imperiled) (G2, 
NatureServe 2005). 


3. Cuscuta campestris Yuncker [nom. nov], Mem. Torrey Bot. Club 18:138. 1932. 
Based on Cuscuta pentagona Engelm. var. calycina Engelm., Amer. J. Sci. 45:76. 1843 [1845]. 
Cuscuta arvensis Beyr. ex Engelm. var. calycina (Engelm.) Engelm., Trans. Acad. Sci. St. Louis 
1:495. 1859. Lectotype (Yuncker 1921): U.S.A. TEXAS: wet prairies, [no date], Lindheimer 126 (MO). 


Stems 0.3-0.5 mm in diameter, yellow to orange. Inflorescences dense, 


164 BRIT.ORG/SIDA 22(1) 


corymbiform or glomerulate of (3-)6-25(-30) subsessile to short pedicellate 
flowers; pedicels 0.3-2.5(-3.5) mm; bracts | at the base of clusters and O-1 at the 
base of pedicels, like those of C. pentagona. Flowers (4-)5-merous, (1.9-)2.1-3.6 
mm (Fig. Ic), membranous, white-creamy when fresh, creamy or golden-yel- 
low when dried; papillae absent; laticifers evident in the calyx and less obvious 
in the corolla and ovary/capsule, isolated or arranged in rows, rotund, ovoid or 
elongated; calyx yellow, reticulate, shiny, rarely obscurely angled, cupulate, about 
as long as corolla tube, divided 2/5-3/5 the length; lobes overlapping but not 
forming or rarely forming obscure angles at sinuses; ovate triangular, not cari- 
nate, margins entire, apex obtuse to rounded (Fig. la); corolla persistent; tube 
campanulate, not saccate, (1.1-)1.5-1.9 mm; lobes spreading, triangular-lan- 
ceolate, (1.1-)1.4-175 mm long, margins entire, apex acute to acuminate, inflexed; 
stamens exserted, shorter than corolla lobes, anthers broadly elliptic, (0.3-)0.4- 
0.5 x 0.25-0.3 mm, filaments 0.4-0.7 mm long; pollen grains 18.5-28 um long; 
tectum perforatum but sexine often persistent in puncta aperture; maximum 
diameter of puncta 0.4 um; supratectal process granular, often in groups of 2-4 
(Fig. 3f); infrastaminal scales equaling or slightly exceeding corolla tube, ob- 
long-ovate to spathulate, rounded, uniformly dense fimbriate; styles evenly fili- 
form, 0.8-16 mm, ca. as long as the ovary; stigmas capitate, globose. Capsules 
indehiscent to irregularly dehiscent, globose-depressed to depressed, 1.3-2.8 x 
1.9-3.8 mm; not thickened or raised around the large interstylar aperture, some- 
times translucent, persistent corolla enveloping 1/3 or less of the capsule base. 
Seeds 4 per capsule, angled, subrotund to broadly elliptic, 112-154 x 0.9-L1 
mm, seed coat cells alveolate/papillate, 32-40 um in diameter; epicuticular wax 
absent; hilum region subterminal, round, 0.45-0.5 mm in diameter, vascular 
scar, 0.085-0.01 mm long, vertical or slightly oblique (Fig. 4 a,b). 2n = 56 
(Fogelberg 1938; Ward 1984). 

Distribution and ecology.—CANADA: Alberta, British Columbia, 
Saskatchewan, Manitoba, Nova Scotia, Ontario and Québec. Darbyshire (2003) 
also mentioned it from NewFoundland, New Brunswick and Prince Edward 
Island. Although we found no herbarium collections, its presence in these prov- 
inces is possible. U.S.A.: Arizona, Arkansas, California, Colorado, Florida, Geor- 
gia, Idaho, Illinois, Indiana, lowa?, Kansas, Kentucky, Louisiana, Massachusetts, 
Maryland, Michigan, Minnesota, Mississippi, Missouri, Montana?, Nebraska, 
Nevada, New Mexico, New Jersey, New York, North Carolina, North Dakota, Ohio, 
Oklahoma, Oregon, Pennsylvania, South Dakota, Tennessee, Texas, Utah, Vir- 
ginia, Washington, West Virginia, Wisconsin, Wyoming. Cuscutd campestris is 
the second most common species in North America, after C. gronovii (Yuncker 
1932), and perhaps the most successful and widespread Cuscuta weed world- 
wide, recorded from South America, Europe, Asia, Africa, and Australia (Holm 
et al. 1997). Flowering Jun-Nov. Hosts: numerous species from hundreds of gen- 
era in various families, including Hydrophyllaceae, Chenopodiaceae, 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 165 


Verbenaceae, Convolvulaceae, Acanthaceae, Brassi Euvhorbi Care 
aceae, Urticaceae, Polygonaceae, Fabaceae, Asteraceae, and others (Gaertner 
1950), growing in a wide variety of climates and soil types; reported not to sur- 
vive on Amaranthus retroflexus, Arctium lappa, Atriplex spp., Brassica nigra, 
Equisetum arvense, Glycine max, selected cultivars of Lycopersicon esculentum 
Portulaca oleracea, Tanacetum vulgare, Vicia villosa, Cyperaceae and Poaceae 
(Gaertner 1950; Parker & Riches 1993; Dawsonet al. 1994; Costea & Tardif 2006). 

Conservation status.—G5 (common) (the same assessment in NatureServe 
2005). 


— 


4. Cuscuta glabrior (Engelm.) Yuncker, Mem. Torrey Bot. Club 18:140. 1932. Cuscuta 
pentagona Engelm. var. ae brior (Engelm.) Gandhi, Thomas, & Hatch, Sida 12:372. 1987. 
Cuscuta verrucosa Engelm. var. glabrior Engelm., Amer. J. Sci. 43:341. 1842. LECTOTYPE (Yucker 
1932): U.S.A. TEXAS. [Harris Co.?|: dry sterile prairies west of Houston, Lindheimer s.n. ( 

The protologue noted that var. glabrior [was growing] “with the preceding variety [ie., C 
verrucosa var. hispidulal, F Lindheimer: on Petalostemon, Drummond (Grd collection, No. 

247).” Choice of the Drummond collection (III, 247) as lectotype of Cuscuta verrucosa Engelm. 
var. glabrior has been associated with Yuncker (1921), but there Yuncker instead chose the 
DEdnond type for Cpe ntagona Var. VErrucosd, citing C. verrucosa var. glabr iorasa synonym. 


Cuscuta verrucosa Engelm., Amer. J. Sci. 43:341. 1842 oo. invalid, non Sweet 1823). Cuscuta 
rvensis var. verrucosa Engelm., Trans. Acad. Sci. St. s 1:495. 1859. Cuscuta pentagona 
Engelm. var. verrucosa (Engelm.) Yuncker, Illinois it Monogr. 6:142. 1921. LECTOTYPE 
(Yuncker 1921): U.S.A. TEXAS: no other locality, Drummond III 247 (MO; ISOLECTOTYPE: TEX). 
Engelmann described C. verrucosa and two varieties of it: var. hispidula (treated by us as C. 
indecora var. indecora) and var. glabrior. He cited calectoe associated with both varieties 
but none for C. verrucosa in its typical form. In 1859, in valid use of the epithet as C. arvensis 
var. verrucosa, he cited “Texas, Drummond! III. 247; Lindheimer! 127” and others as syntypes. 
Yuncker (1921, p. 52), under the combination C. pentagona var. verrucosa, indicated that 
“Drummond III 247” at MO was the type (lectotype). 

Cuscuta arvensis Beyr. ex Engelm. var. pubescens Engelm., Trans. Acad. Sci. St. Louis 1:495. 1859. 
Cuscuta pentagona Engelm. var. pubescens (Engelm.) Yuncker, Illinois Biol. Monogr. 6:142.1921. 
Cuscuta glabrior var. pubescens (Engelm.) Yuncker, Mem. Torrey Bot. Club 18:141. 1932. LEc- 
TOTYPE (Yuncker 1921): “Texas (... L ica in ee )’? (MO). Engelmann’s protologue cited 
“Western Texas, Lindheimer! Wright! 135 (57 


Stems 0.35-0.5 mm in diameter, creamy. Inflorescences loose to compact, 
glomerulate or corymbiform of 3-25(-30) subsessile to pedicellate flowers; 
pedicels 0.8-4(-5) mm; bracts like in C. campestris. Flowers 5-merous, |.4-2.1(- 
2.5) mm (Fig. 2a), membranous, white or commonly reddish when fresh, yel- 
low to reddish-brown when dried; papillae present on the perianth, ovaries/ 
capsules and sometimes on the pedicels (Fig. 2b); laticifers obvious in the calyx 
and less evident in the corolla and ovary /capsule, isolated or arranged in rows, 
rotund, ovoid or elongated; calyx yellow to reddish-brown, shiny, + reticulate, 
cupulate, about as long as corolla tube; divided 1/2-2/3, lobes not overlapping, 
ovate triangular, not carinate, margins entire, apex obtuse to subacute; corolla 
persistent; tube campanulate-globose, saccate between the lines of stamen at- 
tachments, (1.1-)1.5-2.1 mm long; lobes spreading to reflexed, lobes 1.2-1.75mm 


166 BRIT.ORG/SIDA 22(1) 


long, triangular to sublanceolate, margins entire, apex acute to acuminate, in- 
flexed; stamens exserted, shorter than corolla lobes, anthers broadly elliptic to 
elliptic-oblong, 0.44-0.67 X 0.22-0.3 mm, filaments 0.4-0.7 mm; pollen grains 
18-22.6 um long; tectum imperforatum or witha few puncta; maximum diam- 
eter of puncta 0.2 1m; supratectal process granular to conical + uniformly dis- 
tributed; infrastaminal scales reaching the filament bases, ovate to spatulate, 
rounded, uniformly dense fimibriate; styles evenly filiform, 0.9-1.6 mm, as long 
as or longer than the ovary; stigmas capitate, globose. Capsules indehiscent to 
irregularly dehiscent, globose-depressed to depressed, 1.5-2.8 x 2.1-3.5mm, not 
thickened or raised around the medium to large interstylar aperture, not trans- 
lucent, persistent corolla enveloping 1/2-2/3 of the capsule. Seeds 4 per cap- 
sule, angled, subrotund to broadly elliptic, 0.95-L15 x 0.85-1 mm, seed coat cells 
alveolate/papillate, 35-50 um in diameter (Fig. 4c), epicuticular wax absent; 
hilum region subterminal, round, 0.13-0.2 mm in diameter, vascular scar 0.03- 
0.05 mm long, oblique. 2n = ? 

Distribution and ecology.—U.S.A.: Louisiana, New Mexico, Oklahoma, 
Texas, Utah; Mexico. Flowering Jun-Sep. Hosts: wide variety of herbaceous spe- 
cies, including Amaranthus, Ambrosia, Amphiachyris, Asclepias, Convolvulus, 
Coreopsis, Croton, Dalea, Dyschoriste, Evolvulus, Gaura, Gilia, Hedeoma, 
Helenium, Lespedeza, Liatris, Machaeranthera, Medicago, Mimosa, Plantago, 
Prosopis, Tragia, Oenothera, Justicia, Polygonum, Ruellia, Solanum, 
Symphyotrichum, Thelesperma, Verbena. 

Conservation status.-G4G5 (the same assessment in NatureServe 2005). 

According to Yuncker (1965, p. 6), Cuscuta pentagona var. pubescens 
(Engelm.) Yuncker “represents the extreme degree of papillation which covers 
the flower and extends down onto the pedicels. In var. glabrior the ovary and 
capsules are almost always more or less papillate while the corolla shows con- 
siderable variation in this character.” We find continuous variation, and two 
varieties cannot be separated. Occasional plants that are hardly papillate can 
be distinguished from C. campestris by their calyx lobes, which are broadly 
triangular-lanceolate and not overlapping at the base and by the corolla envel- 
oping the mature capsule (Fig. 2a). 


~— 


5. Cuscuta runyonii Yuncker, Bull. Torrey Bot. Club 69:541. 1942. Type: U.S.A. TEXAS. 
Hidalgo Co.: La Joya, occasional, on dry hill tops, clay, 45 m, flowers cream white, hosts: 
Coldenia canescens and Nama hispida, 8 Jun 1941, Runyon 2732 (HOLOTYPE: US; ISOTYPES: F, 
GH, MO, NY, TEX). 

Stems 0.35-0.5 mm in diameter, yellow-orange. Inflorescences loose, + umbel- 

late of 2-15(-25) subsessile to pedicellate flowers; pedicels 0.8-4(-5) mm; bracts 

as in C. campestris. Flowers 5-merous, 2.5-3.5(-4) mm (Fig. 2c), white to creamy 
when fresh, reddish-brown when dried, membranous, papillae present on the 
perianth, ovary/capsules and sometimes on the pedicels; laticifers evident in 


—_— 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 167 


the perianth and less obvious in the ovary/capsule, isolated or arranged in rows, 
rotund, ovoid or tangentially elongated; calyx reddish-brown, + reticulate, shiny, 
cupulate, shorter than corolla tube or about equaling it, divided 1/2-2/3 the 
length, lobes not basally overlapping, triangular, sometimes carinate, base of 
calyx corresponding to each lobe with a reflexed spur-like projection, 0.1-0.6 
mim long (Fig. 2c), and often with smaller protuberances on the midveins of 
calyx lobes, margins entire, apex obtuse to subacute; corolla persistent; tube 
campanulate-globose, saccate between the lines of stamen attachments, (1.2- 
)1.6-2.3 mm, lobes reflexed, 1.2-2 mm, triangular-ovate to lanceolate, margins 
entire, apex acute to acuminate, inflexed; stamens exerted, shorter than corolla 
lobes anthers broadly elliptic to elliptic-oblong, 0.4-0.6 x 0.25-0.3 mm, fila- 
ments 0.4-0.6 mm long; pollen grains 18-23.8 ,1m long; tectum imperforatum 
or with a few puncta; maximum diameter of puncta = 0.3 um; supratectal pro- 
cess granular to conical + uniformly distributed; infrastaminal scales reach- 
ing the filament bases, oblong to spatulate, rounded, uniformly dense fimbri- 
ate; styles evenly filiform, 0.9-1.8 mm, equaling to longer than the ovary; stigmas 
capitate, globose. Capsules indehiscent to irregularly dehiscent, globose-de- 
pressed, 1.6-3 X 2-3.4 mm; not thickened or raised around the large interstylar 
aperture large, not translucent, persistent corolla enveloping 1/2-1/lof the cap- 
sule. Seeds 4 per capsule, angled, ovate to broadly elliptic, 0.95-1.40 x 0.85-L1 
mm, seed coat cells alveolate/papillate, 30-50 um in diameter; epicuticular wax 
present with longitudinal; rodlets hilum region subterminal round to subrotund, 
0.30-0.40 mm, vascular scar, 0.08-0.09 mm long, oblique (Fig. +d). 2n =? 

Distribution and ecology.—U.S.A.: Texas. Flowering Mar-Apr(-May, -July, - 
Oct). Hosts: species of Dalea, Dyschoriste, Erigeron, Gutierrezia, Hymenoxys, 
Justicia, Linum, Melampodium, Oenothera, Nama, Spermolepis, Tetraneuris, 
Tiquilia, Thamnosma, Thelesperma. 

Conservation status.—G3 (vulnerable) (G4, NatureServe 2005). 


6. Cuscuta plattensis A. Nels. Bull. Torrey Bot. Club 26:131. 1899. Type: U.S.A. WYOMING. 
[Carbon Co.?| Platte Canyon, 27 Aug 1896, A. Nelson 2768 (HOLOTYPE: RM; IsOTYPEs: NY, US). 


Stems 0.15-0.3 mm in diameter, yellow to pale orange. Inflorescences loose, 
paniculiform of 2-10(-15) subsessile to pedicellate flowers; pedicels 0.5-2.5(-3) 
mm.; bracts asin C. campestris. Flowers 5-merous, 3-4.6(-5) mm (Fig. 2d), mem- 
branous, white-creamy when fresh, whitish-yellow when dried; papillae ab- 
sent; laticifers present in the calyx and less obvious in the corolla and capsule, 
isolated, ovoid to elongated; calyx yellow, reticulate, not shiny, cylindric-cupu- 
late, ca. equaling the corolla tube, divided 1/2-2/3, lobes slightly basally over- 
lapping, ovate triangular, not carinate, margins entire, obtuse to subacute; co- 
rolla persistent; tube narrowly campanulate to cylindric campanulate, not 
saccate, 1.9-2.5 mm, lobes spreading to reflexed, 1.4-2.2 mm long, triangular, 


168 BRIT.ORG/SIDA 22(1) 


margins entire, apex narrowly acute, inflexed; stamens exserted, shorter than 
corolla lobes anthers subrotund, broadly elliptic to elliptic-oblong 0.4-0.7 x 
0.3-0.4 mm, filaments 0.3-0.4 mm; pollen grains 18-26.3 um long: tectum 
imperforatum or with a few puncta; diameter of puncta 0.2 pm; supratectal 
process granular + uniformly distributed (Fig. 3c); infrastaminal scales 3/4- 
4/5 of the corolla tube, oblong-spathulate, rounded uniformly dense fringed to 
fimbriate; styles uniformly filiform, 1.3-L6 mm, about as long as the ovary; stig- 
mas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose to 
globose-depressed, 1.8-3.2 x 2.2-3.6 mm, not thickened or raised around the 
large intersylar aperture; not translucent, withered corolla persistent envelop- 
ing 1/2 or more of the capsule base. Seeds 1-4 per capsule, dorsoventrally com- 
pressed, broadly elliptic to obovate, 1.07-1.42 x 0.9-1.21 mm, seed coat cells al- 
veolate/papillate, 15-20 1m in diameter; epicuticular wax absent; hilum region 
subterminal, subrotund, 0.30-0.35 X 0.28-0.30 mm. vascular scar 0.20-0.25 mm 
long, vertical to slightly oblique (Fig. 4f). 2n = ? 

Distribution and ecology.—U.S.A.: Nebraska, Washington, and Wyoming. 
Flowering Aug. Hosts: species of Grindelia, Helianthus, Humulus Psoralea, Ru- 
bus, Solidago. 

Conservation status.—Gl (critically imperiled) (the same assessment in 
NatureServe 2005). 


7. Cuscuta sandwichiana Choisy, Mem. Soc. Phys. Hist. Nat. Geneve 9:280. 1841. 
Tyre: U.S.A. HAWAII Sandwich Islands, 1830, Gaudichaud-Beaupré I (HOLOTYPE: G-DC, frag- 
ment NY; ISOTYPE: P, fragment NY). 


Stems, 0.2-0.5 mm, yellow-orange. Inflorescence loose, paniculiform of 3-7(-1D 
pedicellate flowers; pedicels 2-5 mm; bracts | at the base of clusters, usually 
lacking at the base of pedicels, membranous, narrow-triangular, I-1.6 x 0.7-1 
mm, margins entire, apex acute. Flowers (4-)5-merous, 3-4(-5) mm long (Fig. 
5a), fleshy, white when fresh, yellow-brownish when dried; papillae absent; la- 
ticifers conspicuous in the corolla lobes, sometimes in the calyx and ovary/ 
capsule as well, isolated or in rows, ovoid to elongated; calyx yellow-brown, not 
reticulate or shiny, cupulate, almost as long as corolla tube, divided 1/2-1/3 the 
length, lobes not basally overlapping, triangular-ovate, in some flowers cari- 
nate, margins entire, apex obtuse to subacute; corolla persistent; tube campanu- 
late to globular, not saccate, 2-3 mm long, lobes erect to slightly spreading, 0.8- 
L5 mm long, ovate-triangular, margins entire, apex obtuse to subacute with 
inflexed tips; stamens mostly included, shorter than corolla lobes, anthers 
broadly-elliptic 0.5-0.7 x 0.4-0.5 mm, filaments 0.2-0.7 mm; pollen grains 22- 
27.5 um long; tectum imperforatum or with a few puncta; maximum diameter 
of puncta 0.3 um; supratectal process granular to conical, + uniformly distrib- 
uted; infrastaminal scales absent or reduced, bifid, truncated or triangular, dis- 
tally with a few teeth; styles uniformly stoutish, 0.5-1L1 mm, ca. as long as the 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 169 


ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, 
globose to globose-depressed, 2.5-3.5 mm X 33-4 mm, + thickened but not 
raised around the large interstylar aperture, not translucent, withered corolla 
surrounding the capsule. Seeds usually 2 per capsule, angled, elliptic to broadly- 
elliptic, 11-2 x 1-17 mm, seed coat cells alveolate/papillate, 35-50 um in diam- 
eter, epicuticular wax absent; hilum region subterminal, round, 0.30-0.45 mm 
in diameter, vascular scar 0.19-0.25 mm long, vertical or slightly oblique (Fig. 
5b). 2n =? 

Distribution and ecology.—U.S.A.: Hawaii. Flowering summer-fall JQun- 
Oct). Hosts: plants in seashores and arid habitats. 

Conservation status.—G2 (imperiled) (the same assessment in NatureServe 
2005). 


8. Cuscuta obtusiflora Kunth var. glandulosa Engelm., Trans. Acad. Sci. St. Louis 

Q?. 1859. Cuscuta gla ndulosad (Engelm.) Small, Fl. S.E. U.S. 969, 1337. 1903. LECTOTYPE 

(Yuncker 1921): U.S.A. GEORGIA. [Muscogee Co.]|: Columbus, 1838, Boykin s.n. (MO). The 
protologue cited numerous collections in addition to the Boykin one. 


Stems 0.25-0.4 mm in diameter, orange. Inflorescences dense, glomerulate of 
5-18 sessile or subsessile flowers; pedicels absent to | mm; bracts 1 at the base of 
clusters and 0-1 at the base of pedicels/flowers, membranous, ovate, 0.5-1.3 x 
0.2-1.1 mm, margins entire, apex acute to obtuse. Flowers 5-merous, 1.8-2.5 mm, 
membranous, creamy white when fresh, yellow-brown when dried; papillae 
absent; laticifers numerous, evident in the perianth and ovary/capsule, isolated, 
round to ovoid; ealyx yellow-brownish not reticulate or shiny, shallowly cupu- 
late, about equaling the corolla tube, divided ca. 1/2 the length, lobes barely 
basally overlapping, ovate, not carinate, margins entire, apex obtuse; corolla 
persistent; tube campanulate, not saccate, 1-15 mm, lobes erect to spreading, 
0.8-1.5 mm long, ovate to ovate-oblong, margins entire, apex obtuse, straight; 
stamens exserted, shorter than corolla lobes, anthers ovate to broadly elliptic, 
0.3-0.4 x 0.2-0.3 mm, filaments 0.4-0.6 mm; pollen grains (18-)20-24(-27) um, 
tectum perforatum; maximum diameter of puncta 0.6 um; supratectal process 
granular to conical + uniformly distributed (Fig. 5c); infrastaminal scales reach- 
ing the filament bases, oblong, rounded, with a few basal fimbriae, densely 
fringed in the distal 3/4; styles uniformly stoutish, 0.4-1.1 mm, shorter than or 
equaling the ovary, stigmas capitate, globose. Capsules indehiscent to irregu- 
larly dehiscent, depressed-globose, 1.5-3 x 2.5-4 mm, not thickened or raised 
around the large interstylar aperture, not translucent, withered corolla persis- 
tent at the base. Seeds usually 4 per capsule, dorsoventrally compressed, broadly- 
ovate to broadly elliptic, 14-156 x 1.23-1.35 mm, seed coat cells alveolate/papil- 
late, 20-30 pm in diameter, epicuticular wax usually present (Fig. 5d). 2n = ? 
Distribution and ecology.—U.S.A.: Alabama, Arkansas, Connecticut, Florida, 
Georgia, Kentucky, Louisiana, Minnesota, Mississippi, New York., Oklahoma., 


170 BRIT.ORG/SIDA 22(1) 


howe 
=& 
"es 


A 
as 
pk 
ft 
a 


eet Ch Bh 
AS Sey 
Denes ‘3 


‘ et 
ee) ie 
rere 

sf 


— 


Fic. 5. Cuscuta sandwichiana. a. Flower (scale bar = 1 mm);b. seed (scale bar = 0.6 mm). Cuscuta obtusiflora var. glandulosa. 
c. Pollen (scale bar = 6 im); d. seed (scale bar = 0.5 mm). Cuscuta polygonorum.e. Seed—general morphology (scale 
bar = 0.60 mm); f. seed—surface micromorphology (scale bar = 100 Lm). 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 171 


Pennsylvania, Texas; Mexico, West Indies (Puerto Rico, Cuba). Flowering Jul- 
Sep, ~Oct. Hosts: species of Alternanthera, Dalea, Hygrophila, Lythrum, 
Polygonum, Xanthium. 

Conservation status.—G3G4 (vulnerable to apparently secure) (G4, 
NatureServe 2005). 


9, Cuscuta polygonorum Engelm., Amer. J. Sci. Arts 43:342. 1842. Type US.A. Mis- 
souRE W of St. Louis, “bottoms of sink-holes, or margins of ponds”], on Polygonum, Aug 1839, 
Lindheimer s.n. (HOLOTYPE: MO). 


d ,glomerulate 


Stems 0.2-0.4 mm in diameter, yellow-orange. Infl 
of 4-18 sessile to subsessile flowers; bracts 1 at the base of clusters and 0-1 at 
the base of pedicels, membranous, ovate-triangular to lanceolate, 0.3-1 x 0.2- 
0.9 mm, margins entire, apex acute; pedicels absent to | mm. Flowers (3-)4- 
merous, 2-2.7 mm, membranous, white when fresh, yellow-brown when dried; 
papillae absent; laticifers evident in the corolla and the ovary/capsule, isolated, 
round to ovoid; calyx yellow-brownish, not shiny or reticulate, cupulate, ca. 
equaling the corolla tube, divided 1/2-2/3 to the base, lobes not basally over- 
lapping, triangular-ovate, not carinate, margins entire, apex obtuse to rounded; 
corolla persistent, tube cupulate to shallowly campanulate, hot saccate, l-14 
mm, lobes erect, l.1-1L6 mm long, triangular, margin ntire, apex acute, straight; 
stamens exserted, shorter to nearly equaling corel lobes, anthers ovate to 
broadly elliptic, 0.3-0.4 x 0.2-0.3 mm, filaments 0.4-0.7 mm; pollen grains (18-) 
21-24.5 (-26) um long, tectum imperforatum or with a few puncta; maximum 
diameter of puncta 0.3 um; supratectal process granular to conical + uniformly 
distributed; infrastaminal scales usually reaching the filament bases, oblong, 
mostly shallowly bifid with 1-3 fimbriae on each side of the filament attach- 
ment or with a few irregular fimbriae at the apex; styles evenly subulate, 0.4- 
0.9 mm, shorter than the ovary; stigmas capitate, globose. Capsules indehis- 
cent to irregularly dehiscent, depressed-globose, often appearing angled, 1.6-3 
< 2.5-5 mm, not thickened or raised around the large interstylar aperture, not 
translucent, withered corolla persistent the capsule base. Seeds usually 4 per 
capsule, dorsoventrally compressed, subrotund, broadly ovate to broadly el- 
liptic, 145-16 x 1.25-1.39 mm, seed coat cells alveolate/papillate or poliedric, 
20-30 um in diameter, epicuticular wax usually present; hilum region subter- 
minal scar area slightly raised, elliptic, 0.45-0.6 x 0.3-0.35 mm; vascular scar 
0.15-0.2 mm, vertical or slightly oblique (Fig. 5e,f). 2n = 

Distribution and ecology.—CANADA: Ontario, Québec. U.S.A.: Arkansas, 
Connecticut, District of Columbia, Delaware, Iowa, Illinois, Indiana, Kansas, 
Kentucky, Louisiana, Massachusetts, Maryland, Maine, Michigan, Minnesota, 
Missouri, North Dakota, Nebraska, New Jersey, New York, Ohio, Oklahoma, 
Pennsylvania, Rhode Island, Tennessee, Texas, Virginia, Wisconsin. Flowering 


rt 
—" 


172 BRIT.ORG/SIDA 22(1) 


Jul-Oct. Hosts: usually Polygonum but occasionally also on Impatiens, Ipomoea, 
Lycopus, Penthorum, Xanthium, and others. 
Conservation status.—G3 (vulnerable) (G5, NatureServe 2005). 


APPENDIX lL. 
VOUCHERS FOR THE SEM STUDY 


Vouchers are from NY unless otherwise indicated. 


1. Cuscuta pentagona (16 collections examined).—U.S.A. DISTRICT OF COLUMBIA. Eastern 
sae near Mt. Hamilton, 2 Aug 1919, Killip 6338. FLORIDA. Lake Co.: 3 mi S of Sorrento, Welch 
GEORGIA. Whitfield Co.: field E of Dicks Ridge, 1000 ft, 27 Jul 1900, Wilson s.n. INDIANA 
eee Co.: 2 mi W of Bedford, flood plain of Salt Creek, 23 Aug 1934, Kriebel 2518. KANSA S. 
Trego Co.: 19 mi S and 2 mi W of Collier, 6 Aug 1952, David & Harr 4136. MASSACHUSETTS. 
Middlesex Co.: Winchester, Winter Pond, 22 Sep 1908, Fernald & Weatherby s.n. MICHIGAN. 
Kalamazoo Co.: Fort Custer, 12 Aug 1945, Hanes 4541.MISSOURI. Phelps Co.: Jerome, 27 Jul 1931, 
Kellog s.n. NEW JERSEY. Ocean Co.: Bay Head, 31 Jul 1910, Mackenzie 4742.NEW YORK. (no county 
given).Long Island, 8 Aug 1903, Bicknell s.n. NORTH CAROLINA. Franklin Co.: Franklin, 23 Jul 1888, 
Johnson s.n. NORTH DAKOTA. Barnes Co.: Eckelson, 15 Sep 1937, Stevens s.n. Benson Co.: Cran- 
berry Lake, 8 Sep 1930, Stevens s.n. TENNESSEE. Wayne Co.: 4 mi W of Waynesboro, 10 Aug 1934, 
Welch 1382. VIRGINIA. Arlington Co.: 1/4 mi SE Hatfield,9 Jul 1939, Hermann 10391;W of Williamsburg, 
15 Aug 1921, Weatherby 4230 
2.Cuscuta campestris (23 Akai alunt onaiecinalac QUEBEC. paiies te 940, 
Cartier s.n. oNTAR 10, Waterloo Co.: Galt, 26 Sep 1908, Herriot s.n. Wellington Co.: 3 Aug 
1943, Howitt s.n. I rt cel ae Jonesboro, 260 ft,6 Jun 1948, ens 26643, 
Garland Co.: Hot Springs, 500 ft, 17 Jul 1938, Demaree 17932. CALIFORNIA. Los Angeles Co.: Los 
Angeles, 10 Jul 1933, et 10533. IDAHO. Ada Co.: Harrison dee on NE side of Boise, T4N R2E 
Sec 34 SENW, 2710 ft, 1 g 1980, Ertter & Strachon 3951. FLORIDA. Dade Co.: at 14000 SW 8th 
Road, Miami, 12 Nov cen Y).GEORGIA. Clarke Co.: He at 285 Hillcrest, 800 ft, 11 
Nov 1946, Cronquist 4209. INDIANA. Harrison Co.: 4 mi SE of Central, 26 Aug 1945 Deams.n.. Wells 
Co.: 5 mi NE of Bluffton, 26 Aug 1940, Deam 59866. KENTUCKY. Fayette Co.: Gentry-Thompson 
stockyards, Lexington, 2 Sep 1944, McFarland s.n. MARYLAND. Wicomico Co.: Willards,4 Sep 1942, 
Moldenke 13847. NEVADA. Douglas Co.: eae River, ca. 7 mi N of Minden, 4700 ft, 14 Sep 1969, 


gla 
Howell ioe alae EXICO. Grant Co.: | 881, Rusby 295. NEW YORK. Onondaga Co:: Syra- 
cuse, shore of Onondaga a5 cere One GON. Marion Co.: Salem, Willamette River, 13 Jul 
1922, ieee TENNESSE N of Knox lle Wallace’s Orchard, 6 Aug 1943, Wilson 3007. 


TEXAS. Childress Co.: 3.4 mi : a ress,S of Hwy. 287,6 Sep 1945, Whitehouse 10715.Tom Green 
Co.: South Caucho River, a mi N 2 E of Christoval, 29 Aug 1943, Cory s.n. UTAH. Sanpete Co.: 1145, 
R2E,SEC 22,4 mi NE of F a non U-132,9000 ft, 19 Jul 1977, Neese & White 3682. VIRGINIA. 
Nansemond Co.: Suffolk, 24. jul 1893, Heller s.n. 

ta glabrior (15 collections examined).—U.S.A. OKLAHOMA. Murray Co.: Sulphur, 
Platt an Park, 28 May 1 bee Merril 522.Pontotoc Co.: 2 mi SW of Ada,near State Hwy. 12,26 Jun 
1947, Robbins 2607.NEW MEXICO.(no locality given) 1851-1852, Wright 1635. TEXAS. Bell Co.: Near 
Little River, 11 June 1930, ee Killen, 14 Aug 1931, Wolff 3270. Bexar Co.: 16 mi E of San Anto- 
nio, 27 Apr 1921, Schulz 453;San Antonio, 17 Apr 191 oo s.n. Burnet Co.: along State Hwy. 29, 

mi E of Burnet, Webster & Rogers 6476.Coleman Co.: 7 mi E of ee Anna, be Jun 1958, Correll & 

Johnston 19019. Dallas Co.: Dallas, 30 Jun 1872, Hall s.n. Edwards Co.: 29 mi NW of Rocksprings, 27 
Jun 1943, Cory s.n. Ellie Co.: above Red Oak Creek off Waco road, | ; ite Deni 9353. 
Neueces Co.: Corpus Christi, 0-40 ft, 9-12 Apr 1894, Heller s.n. Randall Co.: Buffalo Lake National 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 173 


Wildlife Refuge, 3 Aug 1975, Higgins 9567. elias a low hills in upper big Canyon, ca. 30 mi N of 
Sanderson toward Sheffield, 2800 ft, Warnock 14 
Cuscuta runyonii (5 collections So eee Duval Co.: of Hyw. 44 NE of 
San Diego, 20 Apr 1949, Lundell 14906. Hidalgo Co.: 2 mi N of La Joy,45 m, 13 Jul 1941, Runyon 2825. 
Starr Co.: Brownsville, 50 m,27 Apr 1941, Runyon 2622. Webb Co.: ca. 21 mi from junction of Hwys. 
81 & 83, along Hwy. 83, 1 Apr 1949, Tharp & York 52-163; 46 mi NE of Laredo, route #59, 17 Jul 1958, 
Correll & Johnston 14906 
5. Cuscuta harperi (5 collections examined).—U.S.A. ALABAMA. Cherokee Co.: Lookout 
Mt. at the edge of gorge of May's Gulf, Harper s.n. De Kalb Co.: Desoto State Park, above Fort Payne, 
in Little River Canyon, 8 Oct 1960, Sherman et al. 27757; Glades, 1200 ft, 13 Oct 1962, Demaree 46295; 
Lookout Mt, Jul 1898, Ruth 473. Etowah Co.: Black Creek just above Noccalula Falls, Lookout Mt., 31 
Aug 1911, Harper 147 
scuta plattensis (2 collections examined).—U.S.A.WYOMING. Goshen Co.: 3 mi NE of 
Torrington, T25N R6QW Sec 31 S1/2 SW 1/4, 4200 ft, 15 Aug 1993, Dorn 5470. Niobrara Co.: near 
Newcastle, 24 Jul 1942, Degener & Peiler 16242. 
7. Cuscuta sandwichiana (3 collections examined).—U.S.A. HAWAII. Hawaii Co.: East Maui, 
Makawao, Kaunauhae, La Perouse Bay, 30 Apr 1978, Sy/ aaa Ka’ena Pt.,29 Nov 1969, 
Char 40; O'ahu, Makapu’u Beach Park, 4 Oct 1960, Fujiwara 
8.Cuscuta obtusiflora var. glandulosa (5 eee examine), —U.S.A.ARKANSAS. Ash- 
ley Co.: Lone Prairie, Prairies, 15 Sep 1940. Demaree 21543. va ae White Co.: in the Tippecanoe 


River, ca. 1 mi below Monticello, 14 Aug 1925, Deam 41990. AHOMA. Haskell Co.: stream ca.5 
mi S of the intersection of Hwy 2 with Hwy 31,22 Jul 36 cha , 3387. TEXAS. Dallas Co.: Bachmans 
Dam, bordering lake, wet area, 1 Sep 1942, Lundell & Lund 717,24 Aug 1942, Lundell 11626. 


scuta polygonorum (7 collections eee ana ARKANSAS. St. Francise Co.: 5 
mi S of Forrest City, 19 Sep 1959, McDaniel 1419. INDIANA. Putnam Co.: 2 mi E of Bainbridge, 20 Oct 
1941, Yuncker 10836. NEBRASKA. Otoe Co.: extreme S part of SE1/4,NE1/4, Sec 25, T8N-R14E, 11 Sep 
1974, Sutherland 4096. NEW JERSEY, Somerset Co.: Millstone, plaine de débordement de la riviére 
Millstone, 4 Aug 1956, Lavoie 452 (QUE). NEW YORK. [county not indicated; Nassau Co.?] Long Island, 
Plattsdale, 19 Sep 1925, Ferguson 4608. OHIO. Ottawa Co.: Duck Club,Winous Pt.,4 Aug 1949, Core & 
Anderson s.n. WISCONSIN. Winnebago Co.: swampy area along the shore of lake Winnebago, N be- 
yond the end of Hazel St., N of Murdock, in Gin 25 Jul 1965, Arriman 598 (UNB) 


ACKNOWLEDGMENTS 


We thank directors and/or f ACAD, ALTA, ARIZ, ASU, BRIT, DAO, 
F GH, HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, 
RSA, SASK SES, TEX GLE TUR UBG UGG | BPS -UNB UNM, US-USAS UWO, 
UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Dan Austin, Lytton 
Musselman and Alan Prather provided useful comments and suggestions for 
an earlier version of the manuscript. Thierry Deroin helped us with informa- 
tion about the type of Cuscuta sandwichianaand kindly donated all the Cuscuta 
fragments annotated by Yuncker (held in P) to WLU. Special appreciation goes 
to NY staff for approving and preparing the numerous voluminous loans. Al- 
exandra Smith assisted us with the scanning electron microscope. 


REFERENCES 


Austin, D.F.1979.Comments on Cuscuta — for collectors and curators. Bull. Torrey Bot.Club 
106:227-228. 


174 BRIT.ORG/SIDA 22(1) 


Austin, D.F. 1986. Cuscutaceae. In:T. Barkley, ed. Flora of the Great Plains. University Press of 
Kansas, Lawrence. Pp. 661-666. 

BartHtort, W,, C. Neinuis, D.Cutcer, F. Dirscy, |. MEUSEL, |. THEISEN, and H.WiLHeLmI. 1998. Classifica- 
tion and terminology of plant epicuticular waxes. Bot. J. Linn. Soc. 126:237-260. 

Beuz, T. 1986. A revision of Cuscuta sect. Cleistogrammica using phenetic and cladistic 
analyses with a comparison of reproductive mechanisms and host preferences in spe- 
cies from California, Mexico, and Central America. Ph.D. diss., Univ. of California, Berkeley 

Buia, A. 1938. Cuscutele Romaniei. Bull. Fac. Agron. Cluj 7:1-147. 

Costes, M.and FJ. Taroir. 2006. Biology of Canadian weeds. Cuscuta campestris Yuncker, C. 
gronovii Willd. ex Schult.,C. umbrosa Beyr.ex Hook.,C. epithymum (L.) Land C.epilinum 
Weihe. Canad. J. Pl. Sci. 86:293-316. 

Costea, M.,G.L.Nesom, and F.Taroir.2005. Taxonomic status of Cuscuta nevadensis and Cuscuta 
veatchii (Convolvulaceae). Brittonia 57:264—272. 

Costea, M., G.L. Nesom, and S. Steranovic, 2006a. Taxonomy of the Cuscuta salina-californica 
complex (Convolvulaceae). Sida 22:177-195. 

Costea, M., G.L. Nesom, and S. Steranovic. 2006b. Taxonomy of Cuscuta gronovii and Cuscuta 
umbrosa (Convolvulaceae). Sida 22:197-20/7. 

Costes, M., G.L. Nesom, and S. Steranovic. 2006c. Taxonomy of the Cuscuta indecora 
(Convolvulaceae) complex in North America. Sida 22:209-225. 

Crupen, RW. 1977. Pollen-ovule ratios: a conservative indicator of breeding systems in 
flowering plants. Evolution 31:32-46. 

Dawson, J.H. 1984.A vegetative character that separates species of Cuscuta. Proc. 3rd Interntl. 
Symposium on Parasitic Weeds. Pp. 184-187. 

Dawson, J.H., L.J.Mussetman, P.Wotswinket, and |. Dorr. 1994. Biology and control of Cuscuta. 
Rev.Weed Sci. 6:265-317. 

Dareysuire, S.J. 2003. Inventory of Canadian agricultural weeds. Agriculture and Agri-Food 
Canada, Research Branch, Ottawa, Ontario. 396 pp. Electronic Publication: <http:// 
res2.agr.ca/ecorc/weeds x herbes/title-titre x e.htm> 

ENceLMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with 
critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 
1:453-523 

Foceteers, 5.0. 1938. The cytology of Cuscuta. Bull. Torrey Bot. Club. 65:631-645. 

GaerINeR, E.E. 1950. Studies of seed germination, seed identification, and host relation- 
ships in Dodders, Cuscuta spp. Mem. Cornell Agric. Exp. Sta. 294:1—56. 

GANDHI, K.N.and R.D.THomas. 1983.Variations in the floral structure of Cuscuta L.Phytologia 
53:184-186. 

GANDHI, K.N., R.D. THomas, and S.L. Hatcu. 1987. Cuscutaceae of Louisiana. Sida 12:361-379. 

Gomez, JM. 1994. Importance of direct and indirect effects in the interaction between a 
parasitic angiosperm (Cuscuta epithymum) and its host (Hormatophylla spinosa).Oikos 
71:97-106 

Hoen, P. 1999. Glossary of pollen and spore terminology. Laboratory of Palaeobotany and 


Att / | 


Palynology, Utrecht. <http://www.bio.uu.nl/~palaeo/glossary/glos-int.Atm> 


COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 175 


Howm, L., J. Dott., E. Houm, J.PANCHO and J. Hersercer. 1997.World weeds:natural histories and 
distribution. John Wiley & Sons Inc., Toronto, ON. 

Kaatesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T. 
and C.A.Meacham. 1999. Synthesis of the North American flora, Version 1.0.North Caro- 
lina Botanical Garden, Chapel Hill. 

Knepper, D.A., R.A. Creacer and L.J. Mussetman. 1990. Identifying dodder seed as contami- 
nants in seed shipments. Seed Sci. Technol. 18:731-741 

Kuur, J. 1969. The biology of parasitic flowering plants. Univ. of California Press, Berkeley. 

LysHebe, B.O. 1984. Seed structure and germination in Cuscuta pedicellata with some notes 
on Cuscuta campestris. Nordic J. Bot. 4:669-664. 

LysHebe, B.O. 1985. Morphological and anatomical features of Cuscuta pedicellata and C. 
campestris. Nordic J.Bot.5:65-77. 

Mussetman, LJ. 1986. The genus Cuscuta in Virginia. Castanea. 51:188-196. 

NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web applicatian) 
Version 4.0. NatureServe, Arlington, Virginia. <http:// .org/explorer> 
(Accessed: November 16, 2005). 

Parker, C.and C.R.RicHes. 1993.Parasitic weeds of the world. Biology and control.CAB Inter- 
national, Wallingford, UK. 

Pazy, B. and U. PLiTMANN 1995. Chromosome divergence in the genus Cuscuta and its sys- 
tematic implications. Caryologia 48:1 73-180. 

Prater, L.A.and R.J.Tyrt. 1993.The biology of Cuscuta attenuata Waterfall. Proc. Okla. Acad. 
SCL 73-13 

Severova, E.E. 1991. Morphology and taxonomy of dodder (Cuscuta L.) of the European 
part ofthe USSR. Byull. Mosk. Obshch. Ispyt, Prir,, Biol. 96:59-63. 

USDA, NRCS. 2004. The PLANTS database. National Plant Data Center, Baton Rouge, LA. 
Electronic Publication,Version 3.5: <http://plants.usda.gov> 

Verocourt, B. 1948. Biological flora of the British Isles. Cuscuta L. J. Ecol. 36:356-365. 

Warp, D.E. 1984. Chromosome counts from New Mexico and Mexico. Phytologia 56:55-60. 

Yuncker, 1.G. 1921. Revision of the North American and West Indian species of Cuscuta. 
Illinois Biol. Monogr. 6:91-231. Reprinted 1970, Johnson Reprint Company, N.Y. 

Yuncker, 1.G. 1932. The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. 

Yuncker, 1.G. 1942. Three new Cuscutas from western North America. Bull. Torrey Bot. Club 
69:541-543. 

Yuncker, T.G. 1943. Nomenclatural changes in the genus Cuscuta and notes on some 
American species. Bull. Torrey Bot. Club 70:61-67. 

Yuncker, T.G. 1965. Cuscuta. North American flora, ser. 2,4:1-51. 


176 BRIT.ORG/SIDA 22(1) 


Book REVIEW 

DieTRICH FROHNE and Haws J. Pranper. Patrick McKinney, Kirk Cumpston (Con- 
sult. Eds.). (Translated by Inge Alford). 2005. Poisonous Plants: A Hand- 
book for Doctors, Pharmacists, Biologists and Veterinarians, Second Edi- 
tion. (ISBN 008192-750-3, hbk.). Timber Press Inc, 133 S.W. Second Avenue, 
Suite +50, Portland, OR 97204-3527, U.S.A.(Orders: wwwtimbet press.com, 
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). 
$150.00, 480 Pp. 224 color photos, 131 b/w illustrations, 20 tables, 115 dia- 
grams, 91/2" x 10 1/2". 


Many seemingly innocuous plants harbor toxins that can cause serious illness and even n. This 


authoritative text is intended for pharmacists, poison control center personnel, doctors, veterinar- 


ians, and other sores who deal with poisoning cases, although it is useful to those interested 
in wild foods and wi iderat Hone medicinal herbs. It epens! with a concise introduction to poisonous 


plants, including hov ning cai yee doi pected cases [ poisoning and descrip- 
k contains clear descripti - poisonous plants, 


tions of toxic ees als. The rest of fig 
arranged by family from Aceraceae to ee ‘The text is aoe with descriptions of 
macroscopic and microscopic plant morphology, symptoms, treatments, and references. Excellent 
color photographs illustrate sae section a met English cauion of a sie was ne 20 
gli -} 


years ago. This second E 
dix with notes on Eaiconetie ee found in Nor th America. Not intended to ae a field ¢ muitde hae 
book is still an excellent reference book.—Marissa Oppel, MS, Herbarium Technician, Botanical Re- 
search Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


Book Notice 
CARMEN ULLOA ULLoA and D.A. Neil. 2005. Cinco aitos de adiciones a la flora del 
Ecuador: 1999-2004. (ISBN 9978-09-578-0, pbk.). Universidad Técnica 
Particula de Loja, Missouri Botanical Garden, Funbotanica. Editorial, UTPL, 
Loja. (Orders: Available at Missouri Botanical Garden Press, http:// 
www.nbgpress.info/). $16.00, [Reduced price availabl« only at UTPL, Loja]. 
ToD o1/ 2 oC AL 
Summary.—Of the 1246 new additions to the Ecuadorian flora, 820 are new taxa described from 
Ecuadorian material, 337 names are new records for Ecuador, and 89 are taxonomic changes. The 
new total for the flora is of 17,058 species of vascular plants. Some 242 new bibliographic references 


Qu 


are liste 


SIDA 22(1):17 176. 2006 


TAXONOMY OF THE CUSCUTA SALINA-CALIFORNICA 
COMPLEX (CONVOIVULACEAE) 


M ihal Coster (corresponding author) Guy |. Nesom 
Department of Biology Botanical Research Institute of Texas 
Wilfrid Laurier University Pecan Stree 
75 University A West Fort Worth, Texas 76102-4060, U.S.A. 
Waterloo, Ontario gnesom@brit.org 


N2L 3C5, CANADA 
meostea@wlu.ca 


Sasa Stefanovic 


Department of Biology 
vaivelsigye oF Toronto at Mississauga 


I UUYU hodad 
Missis issauga, Ontario 
LSL 106, CANADA 
ABSTRACT 


The Cuscuta califor plex (Cuscuta subsect. Californicae) is recircumscribed to include the 


species from aieceee PS ioineticae The species include C. salina, C. susksdorfii, C. californica, C. 
occidentalis, C. subinclusa, and C. howelliana. Asummary of relevant taxonomic information is pro- 


vided, along with a key to species and varieties, synonymy, distribution, host range, and conserva- 
tion status. Morphology and micromorphology of flowers, seeds, and capsules are analyzed and il- 
lustrated. 


RESUMEN 


uscuta subsect. Californicae) Aci i incluir las especies 


El complejo Cuscuta c lifornica ( 
de subsect. Subinclusae. Las especies ee C. salina, C. susksdorfii, C. californica, C. occidentalis, 


C. subinclusa, y C. howelliana. En este trabajo se presenta un resumen de la informacion taxonémica 


junto con una clave para identificar las especies y variedades, sinonimos, distribucion, variabilidad 
de huéspedes y estato de conservacion. Se analizan e ilustran la morfologia y micromorfologia de las 
flores, de y de las capsulas. 


Yuncker (1932) described and defined Cuscuta subsect. Californicae and subsect. 
Subinclusae mainly on the basis of absence or presence, respectively, of 
infrastaminal scales. He suggested that both sections were derived indepen- 
dently but in parallel from subsect. Arvenses. In Yuncker’s view, subsect. 
Californicae comprised five species: C. californica Hook. & Arn., C.occidentalis 
Millsp., C. brachycalyx Yuncker, C.jepsonii Yuncker,and C. sandwichiana Choisy, 
while subsect. Subinclusae included three species: C. salina Engelm., C. 
suksdorfii Yuncker, and C. subinclusa Durand & Hilgard (Yuncker 1932, 1965). 
Cuscuta howelliana Rubtzoff was recently described (Rubtzoff 1966) and noted 
by its author to have similarities to C. suksdorfii of subsect. Subinclusae. 


SIDA 22(1): 177 — 195. 2006 


178 BRIT.ORG/SIDA 22(1) 


Definitions and interrelationships among these taxa are complex. Beliz 
(1993) included Cuscuta brachycalyx, C. occidentalis, and C. suksdorfii as syn- 
onyms of C. californica var. breviflora Engelm. Cuscuta subinclusa and C. 
howelliana were hypothesized to be sister species in a separate clade (Beliz 1986). 
Some taxa were described in one subsection and later were reclassified in the 
other. For example, Engelmann (1859) initially described C. salina as “°C. 
californica var. squamigera” and as C. subinclusa var. abbreviata (see also the 
example below of C. suksdorfii). Cuscuta sandwichiana (Beliz 1986, Costea et al. 
2006a) and C. jepsonii (Beliz 1986; Costea et al. 2006c) do not appear to be evo- 
lutionarily related to this group. 

Circumscriptions of these taxa and an assessment of their we 
are provided here on the basis of the morphology and micromorphology of flow- 
ers, capsules, seeds, and pollen. Conservation status is assessed for all taxa. 


METHODS 


Descriptions of morphology (see Costea et al. 2006a) are based on samples from 
specimens of the NY, JEPS, and UC herbaria (Appendix 1). Measurements and 
pictures were taken with a scanning electron microscope Hitachi S-570 at 15 
KV.Samples were coated with 30 nm gold using an Emitech K 550 sputter coater. 
Conservation status was determined using NatureServe (2005) ranks and 
Criteria. 


TAXONOMY 


Delimitation of the Cuscuta salina-californica complex 

Our observations suggest that the species of subsections Californicae and 
Subinclusae form a single phylogenetic group. We hypothesize that 
infrastaminal scales have undergone a gradual reduction from fimbriate scales 
or ridges in C. salina, to dentate wings in C. suksdorfii, to complete reduction in 
C. californica and C. occidentalis. A similar reduction of infrastaminal scales 
has occurred in C. indecora complex (subsect. Indecorae Yuncker) (Costea et al. 
2006c). Cuscuta subinclusa and C. howelliana have well-developed infrastaminal 
scales and were suggested by Beliz (1986) to form a distinct clade in which a 
cylindric-campanulate corolla has evolved as a specialization to butterfly pol- 
lination. Although C. subinclusa and C. howelliana are distinct in their 
microreticulate pollen (see descriptions below), close similarities in morphol- 
ogy and micromorphology of calyx, corolla lobes, seeds, and capsules (see be- 
low) indicate that they probably belong to the same phylogenetic group as C 
californica, C. salina, and C. suksdorfiti. A cylindric-campanulate corolla some- 
times occurs in C. californica, and Engelmann (1876) noted that “in many re- 
spects [C. salina] is intermediate between the preceding [C. californica] and the 
following species [C. subinclusal.” Yuncker (1932, 1965) observed that the corolla 
of C. subinclusa is “usually showing horizontal ridges between the stamen at- 


— 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 179 


tachments” and we find this characteristic in some flowers of C. californica as 
well. The original description of C. howelliana (Rubtzoff 1966) noted that it is 
similar to C. suksdorfii var. subpedicellata. 

Cuscuta jepsonii and C. sandwichiana were included by Yuncker (1932) in 
subsect. Californicae (the latter species omitted in the treatment from 1965) 
because they exhibit a similar reduction of the infrastaminal scales. Cuscuta 
jepsonii, however, may be evolutionarily related to taxa of Cuscuta indecora com- 
plex (Costea et al. 2006c). Cuscuta sandwichiana is a Hawaiian endemic and 
was hypothesized by Costea et al. (2006a) to belong to the C. pentagona com- 
plex. 

In conclusion, the Cuscuta salina-californica complex in our view includes 
C. salina, C. susksdorfit, C. californica, C. occidentalis, C. subinclusa, and C. 
howelliana, which are recognized here as Cuscuta subsect. Californicae Yuncker 
(including subsect. Subinclusae). Taxonomically and nomenclaturally problem- 
atic species are discussed below. 


Cuscuta subinclusa.—Curran (1885) noted that “from the description, [C. 
ceanothi is]evidently C. subinclusa,’ and the former name has been used by most 
authors (e.g. Yuncker 1965) because it has priority. From Curran’s observation 
it can be inferred that the type collection of C. ceanothi was not available even 
at 1885. This collection may have been destroyed during the fire following the 
San Francisco earthquake from 1906 or even at an earlier date. As observed by 
Beliz 986), it is possible that the name C. ceanothi refers to a species distinct 
from C. subinclusa. The protologue of C. ceanothi states that flowers are urceolate 
with obtuse calyx lobes, features not encountered in C. subinclusa or in any 
species hypothesized to be closely related. In a letter addressed to Engelmann 
on 10 Nov 1860 (Ertter 2003), Behr wrote: “Of Cuscuta I know two kinds, the 


the latter I gave once a diagnosis in the transactions (Proc. Calif.) under the p pres 
liminary name Cuscuta ceanothi. This diagnosis is by the nature of a search 
very incomplete, as for comparison I had only the just-mentioned parasitic kind 
on the Salicornia and none of the more closely related exotics.” In a summary 
of the San Francisco flora (1888), approximately three decades after describing 
C. ceanothi, Behr mentioned C. subinclusa but not C. ceanothi. Without 
neotypification, there apparently is no way to establish the identity of C. 
ceanothi, and until more certainty might exist that C. ceanothi is not a distinct 
and evidently rare species, we use the later name C. subinclusa for the known 
species. 


Cuscuta suksdorfii.— Yuncker (1921) described C. salina var.acuminata Yuncker, 
which he later (1932) treated at specific rank as C. suksdorfii. This species is 
“closely allied with C. salina” from which it differs in the morphology of calyx 
and corolla, infrastaminal scales reduced to dentate wings, and multiseeded 


180 BRIT.ORG/SIDA 22(1) 


capsules. Beliz (1986) originally treated C. suksdorfii as a variety of C. salina, 
but later (1993) she included it as a synonym of C. californica var. breviflora, 
together with C. occidentalis and C. brachycalyx. Indeed, C. suksdorfii shares 
some characteristics with both C. salina (e.g., presence of infrastaminal scales 
as well as seed and pollen morphology—see descriptions) and C. californica/C. 
occidentalis (e.g., multiseeded capsules), but we consider that treating it at spe- 
cific rank better represents the differences in morphology and biology as well 
as the evolutionary relationships between all the taxa involved (see below). 
Cuscuta suksdorfii var. subpedicellata Yuncker with flowers sessile or Peis 
shorter calyx (16-2 mm long) and capsule globose to depressed globose 
treated formally at varietal rank. 


Cuscuta californica and C. occidentalis —By describing C. brachycalyx, Yuncker 
(1932, 1965) obscured the already tenuous distinction between C. californica 
and C. occidentalis. Although homotypic, C. californica var. brachycalyx 
Yuncker and C. brachycalyx have slightly different protologues. Cuscuta 
californica var. brachycalyx has “corolla campanulate, lobes ... shorter than the 
tube...” (Yuncker 1921, p. 62); C. brachycalyx has “corolla campanulate-globose, 
saccate between the stamen attachments, lobes ... shorter than or about equal- 
ing the tube” (Yuncker 1932, p. 159) and “is closely related to C. californica, but 
differs by its very short calyx and more obtuse perianth lobes.” Yuncker did not 
mention the saccate corolla that would separate C. brachycalyx from C. 
californica but that would bring it close to C. occidentalis. Not surprising]y, Beliz 
(1986) concluded that “all names proposed for the numerous perianth and an- 
ther size variants within C. californica do not warrant taxonomic recognition 

. and she included both C. occidentalis and C. brachycalyx as synonyms of C. 
californica var. breviflora (Beliz 1986, 1993). We find that indeed two different 
major entities, corresponding to C. californica and C. occidentalis, can be dis- 
tinguished in most cases, based on a combination of characters. Plants called C. 
brachycalyx by Yuncker, with a short calyx and a long (ca. 2.5 mm) campanu- 
late corolla tube that may become somewhat saccate in fruit, possess all the 
characteristics of C. californica (see below). Although such plants are occasion- 
ally distinct, at other times the calyx/corolla tube ratio may vary even on the 
same plant, from flowers with calyx ca. equaling the corolla tube to flowers 
with calyx ca. 1/2 of the corolla tube length (Fig. 1 a, b,c). Apparently the co- 
rolla tube may continue to grow from the beginning of anthesis until fructifi- 
cation, significantly altering the ratio between calyx and corolla tube lengths. 
Similar infraspecific variation of the ratio between calyx and corolla tube 
lengths may be encountered in C. gronovii (between var. gronovii and var. 
latiflora Engelm.), but here the entities are relatively discrete (Costea et al. 
2006b). For these reasons, C. brachycalyx is here considered conspecific with C. 
californica and not recognized at any rank. 


~— 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 


flowers of the same individual (scale ale m).d talis—flower (scale bar = 
(scale bar = 0.75 mm) ill f C. howelli 7um).g-h ae aS by corolla:g 
C cee h.C. penta (scale bar = 1mm). 


Fic. 1. Morphology of flowers and capsules. a—c. Cuscuta aig var. californica. ae of corolla tube length in 
mm).e. C. suksdorfii — calyx 


182 BRIT.ORG/SIDA 22(1) 


Some plants of this complex produce a short (1.5-2 mm), campanulate-glo- 
bose corolla tube (as in Cuscuta occidentalis) and long anthers/styles (as in C. 
californica), or plants may show reversed character states (corolla tube long 
and campanulate and anthers/styles short). Such intermediate-like plants oc- 
cur at a frequency of ca. 1-2% of specimens examined. It is unclear if they are 
result of hybridization, which has never been clearly documented in Cuscuta, 
or evidence of variation in closely related and incompletely differentiated taxa. 
Because such possibly intermediate plants are relatively few, and to avoid no- 
menclatural changes, we maintain C. californica and C. occidentalis as distinct 
species. Several varieties within C. californica usually can be distinguished, 
based on single characters. Cuscuta occidentalis is less variable; a few collec- 
tions were found to possess papillose flowers like C. californica var. papillosa, 
but the origin of this variation is unclear. 


KEY TO SPECIES OF THE CUSCUTA SALINA~CALIFORNICA COMPLEX 


1. Infrastaminal scales present (Sometimes reduced to ridges), fimbriate 
2. Flowers 5—7(-—9) mm long; calyx ca. 1/2 of the corolla tube, with lobes overlap- 
ping at base; corolla lobes 1/4-1/3 as long as the tube; anthers 0.8-2 mm long 
.C. subinclusa 
2. Flowers 2.8-5(-6.) mm long; calyx ca.equaling or somewhat longer than corolla 
ube, with non-overlapping lobes; corolla lobes + equaling the tube; anthers 
0.3-0.7 mm lon 


3. Flowers 5-merous; calyx and corolla lobes acute to acuminate; capsules ellip- 
tical-ovate, + thickened around the interstylar aperture, with 1 seed 1.C. salina 
3. Flowers 4- and 5-merous; at least some calyx and corolla lobes in the same 
flower long-attenuate; capsules globose to slightly ice 
apically, with 1-4 seeds C. howelliana 
. Infrastaminal scales SOMPISIEY Ansel or represented by lateral, iemiate en 
4. Flowers 4-5 llalobes long ac 
represented by lateral, dentate wings; withered corolla surrounding ae in 
the lower half 4 e suksdorfii 
4. Flowers 5 | lla lot but not ac inf! 


scales completely absent; withered corolla completely enveloping the capsule 
or leaving only its top visi 
5. Flowers short- medic taee corolla not saccate between the stamen attach- 
ments (if slightly saccate then with all the following characteristics); styles 
1.2-3 mm long; anthers oblong to linear,0.7—1.1 mm long; capsule completely 
enclosed by corolla (top not visible); seeds not gg through the corolla 
and pericarp, which are thicker, not semitransparen 5.C. californica 
5. Fl il | il II te bet the st ttachments; 
styles 0.5-1(-1.5) mm,anthers broadly-elliptic,0.25—0.5 mm long; capsule not 
completely enclosed by corolla (at least the top of capsule is visible); seeds 
visible through the thin and semitransparent corolla and pericarp 6.C. 
occidentalis 


1. Cuscuta salina Engelm. in W.H. Brewer, S. Watson, & A. Gray, Bot. Calif. 1:536. 
1876. Grammica salina (Engelm.) Taylor & MacBryde, Canad. J. Bot. 56:186. 1978. LECTOTYPE 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 183 


(Yuncker 1932): U.S.A. UTAH: Rio Virgen, on Suaeda, saline soil, Nov 1885, Remy s.n. (MO, frag- 
ment NY). This was essentially a renaming at specific rank (as “C. salina, pogo n. sp.”) of 
what Engelmann had earlier published as C. subinclusa var. abbreviata and C. californica var. 
squamigera—both varieties were ated in ue y. Yuncker (1932) speci fically referred to 


the MO es Remy, in the herbariur f the Missouri B | Garden”) as the type of 
C.salin 


TInfl eA ae Taser 


Stems ae orange-yellow. : pedicels 
0.5-2.5 mm long; bracts 1(-0), ovate-lanceolate to lanceolate. Flowers 5-merous, 
white, 2.5-5(-6.2) mm long; papillae present or absent in the corolla; laticifers 
conspicuous in the perianth, ovary and capsule. Calyx campanulate to narrow- 
campanulate, about as long as the corolla tube, divided ca. 1/2 the length, glossy 
yellow or brownish when dried, lobes ovate-lanceolate, acute to acuminate, + 
unequal, not basally overlapping or slightly so. Corolla 2.4-4.8(-6) mm long; 
tube cylindric-campanulate to campanulate, 1.2-2.5 mm long, lobes ovate-lan- 
ceolate, acute to acuminate, ca. as long as the tube, erect to spreading, some- 
times basally overlapping. Stamens included or exerted, anthers elliptical, 0.3- 
0.7 mm long, filaments equaling to longer than anthers. Pollen 3(-4)-zonocolpate 
19-22(-26) um, polymorphic, subsphaerical to subprolate, rounded at poles, 
tectum perforatum, puncta, 0.3-0.5 um in diameter, granulate (Fig. 2a). 
Infrastaminal scales oblong, short-fringed, rarely reduced to sparingly fringed 
ridges ca. 1/2-2/3 the corolla tube length. Styles distinct, 0.4-1 mm long; stigmas 
capitate, globose. Capsules elliptical-ovate, 2-3.6 x 1.4-2.1 mm, + thickened 
around the interstylar aperture, indehiscent or irregularly dehiscent, surrounded 
or capped by the withered corolla. Seeds | per capsule, not visible through the 
persistent corolla and pericarp, 1.35-1.57 x 1.25-1.43 mm, + dorsoventrally com- 
pressed, broadly elliptic to subround, hilum subterminal, subround, 0.11-0.14 
x 0.7-0.11 mm, vascular scar linear, 0.02-0.05 mm, oblique; surface of seed coat 
epidermis alveolate when dried and papillate when hydrated, cells 30-40 um 
in diameter. n = 14 (Beliz 1986); 2n = 30 (Pazy & Plitmann 1995). 


KEY TO VARIETIES OF CUSCUTA SALINA 


1. Papillae present on pedicels, calyx and/or corolla;infrastaminal scales represented 


by narrow, sparingly fringed ridges 1c. C. salina var. papillata 
1, Papillae absent; infrastaminal scales narrowly oblong, few-toothed. 

2. Corolla 2-3.5 mm; inland salt flats 1a. C. salina var. salina 

2. Corolla 3-4.8(-6) mm; coastal salt marshes, tidal flats 1b. C. salina var. major 


la. Cuscuta salina var. salina. Cuscuta subinclusa var. abbreviata Engelm., Trans. Acad. Sci. 
St. Louis 1:500. 1859. Type: U.S.A. CALIFORNIA. [Solano Co.]: Mare Island in San Francisco Bay, 
on Arthrocnemum, Wright s.n. HOLOTYPE: MO). 


Cuscuta i Hook. & Arn. var. squamigera Engelm., Trans. Acad. Sci. St. Louis 1499. 1859. 
a salina var. sq uaMugen (Engelm.) Yuncker, Illinois Biol. Monogr. 6, pt. 2-3:71, fig. 126. 

192] Im.) Piper, Contr. U.S. Natl. Herb. 11:455. 1906. Type: U.S.A. UTAH: 

Rio ne ea Sudeds line ol Nov 1855, J. Remy s.n. (HOLOTYPE: P; ISOTYPE: MO, fragment 
NY). Engelmann’s protologue cere noted “J. Remy! in Hb. Mus. Paris.” Yuncker (1921) 
described the type of var. squamigera as “Rio Virgen, (Remy in 1855, ...a fragment in the En- 


BRIT.ORG/SIDA 22(1) 


184 


10 m).b. C. occidentalis (scale bar = 10 m).c.—-d. C. howelliana 


(scale bar = 10 and 3 tm, respectively). e—f. C. subinclusa (scale bar = 10 and 3 |im, respectively). 


{ 


MPs r 


P 


Fic.2.M 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 185 


gelmann Herb.).” 
Distribution and ecology —CANADA: British Columbia. U.S.A.: Arizona, Cali- 
fornia, Nevada, New Mexico, Oregon, Utah, Washing MEXICO: Baja Califor- 
nia. Flowering Apr-Nov. Hosts: Atriplex, Cressa, Centromadia, Jaumea, Plan- 
tago, Salicornia, Salsola, Suaeda, Trichostema, Wislizenia, in inland salt flats, 
marshes, ponds. 
Conservation status.-T4 (apparently secure) (not yet assessed by 
NatureServe 2005). 


1b. Cuscuta salina var. major Yuncker, Illinois Biol. Monogr. 6:161. 1921. Type: U.S.A. 
CALIFORNIA. Santa Clara Co. Palo Alto, frequent on Salicornia in the marshes, 14 Sep 1901, 
Baker 41 (HOLOTYPE: NY; ISOTYPES: CAS, GH, RSA). 
Distribution and ecology.—CANADA: British Columbia. U.S.A.: California, Or- 
egon, Washington. Flowering Jul-Oct. Hosts: various species, especially 
Salicornia, in coastal salt marshes, tidal flats. 
Conservation status.—T3T4 (vulnerable to apparently secure) (not yet as- 
sessed by NatureServe 2005). 


1c. Cuscuta salina var. papillata Yuncker, Bull. Torrey Bot. Club 69: 543. 1942. 
YPE: U.S.A. CALIFORNIA. Mendocino Co.: Fort Bragg, 8-16 Aug 1912, Eastwood 1593 (HOLOTYPE: 
GH, fragment NY). 


Distribution and ecology. —U.S.A.: Arizona, California, Utah. Flowering Jun-Oct. 
Hosts: various species in salt marshes, flats, ponds. 

Conservation status.—T3T4 (vulnerable to apparently secure) (not yet as- 
sessed by NatureServe 2005). 


2. Cuscuta howelliana Rubtzoff, Leafl. W. Bot. 10:335. 1966. Type: U.S.A. CALIFORNIA. 

ke Co. Boggs Lake, open dry margin of the lake, inundated in winter, 20 Aug 1966, Rubtzoff 

& Arnaud 5792 (HOLOTYPE: CAS, ISOTYPES: CAS, GH, OSC, RSA, UC). 

Stems slender, yellow to orange. Inflorescences few- to many-flowered + 
glomerulate cymes, pedicels 0-0.6 mm long; bracts 1-0, lanceolate. Flowers 
embedded in the inflorescence of the host, 4-5-merous, 3-4.5 mm long, whitish, 
papillate-glandular. Papillae present in the calyx and corolla; elongated latici- 
fers present in the calyx and corolla but not obvious (Fig. lf). Calyx campanu- 
late, ca. equaling corolla tube or exceeding it, divided 1/2-2/3 to the base, lobes 
unequal, triangular-ovate, acute, acuminate to long-attenuate and recurved at 
the apex, non-overlapping. Corolla tube cylindrical-campanulate, later more 
or less urceolate, ca. 1.5-2.2 mm long, lobes unequal, ca. equaling the tube, tri- 
eeu as ovate, way pee to nea Aung recurved tips, suberect to spread- 


ing ant 1,0.4-0.7 mm long; filaments 0.1-0.4 mm 
long. Pollen 3(-4)-zonocolpate (15- )17-22(-24) um, polymorph, sphaerical to 
subprolate (subsphaerical ), rounded at poles, tectum perforatum 


to microreticulate, puncta, 0.4-0.8 4m in diameter, granulate (Fig. 2 c, d). 


186 BRIT.ORG/SIDA 22 


= 


1) 


Infrastaminal scales oblong-ovate, fringed, reaching to about the middle of the 
corolla tube; Styles distinct, evenly filiform, 0.4-11 mm; stigmas capitate, glo- 
bose. Capsules globose to slightly depressed, 1.2-1.5 « 0.8-1.2 mm, not thick- 
ened apically, indehiscent or irregularly dehiscent, completely enclosed by the 
withered corolla and latter capped by it. Seeds 1-4 per capsule visible through 
the semi-transparent corolla and pericarp, 0.9-1.43 x 0.8-1.3 mm, dorsoventrally 
compressed to slightly angled, subround to broadly-elliptic, hilum subtermi- 
nal, scar area clearly differentiated from the rest of the seed, broadly elliptic 
0.21-0.25 x 0.12-0.15 mm, hilum linear 0.05-0.06 mm, vertical; heterogeneous; 
some areas are alveolate with cells 35-50 um in diameter; some areas irregu- 
larly wrinkled (Fig. 3c, e, f, g). 2n = 26 (Beliz 1986). 

Distribution and ecology.—U.S.A.: Endemic in California. Flowering Aug- 
Sep. Hosts: mostly Eryngium aristulatum, E. vaseyi, E.castrense, E.alismaefolium, 
Navarretia leucocephala, and N. minima, but also on Polygonum kelloggii, 
Epilobium pygmaeum, margins of vernal pools. 

Conservation status.—G2G3 (imperiled to vulnerable) (G3, NatureServe 
2005). 

The biology of this species deserves future study. The parasite becomes 
strictly localized to the inflorescence region of the host. Flowers of C. howelliana 
develop inside the dense host inflorescences and apparently synchronize their 
anthesis with that of the host's flowers. The parasite achieves both protection 
and proximity to the flux of assimilates intended for the development of host’s 
reproductive structures. 


— 


3. Cuscuta subinclusa Durand & Hilgard, J. Acad. Nat. Sci. Philad. ser. 2, 3:42. 
Type: U.S.A. CALIFORNIA. [Los Angeles Co]: Tejon Pass, on a willow, Heermann s.n. CHO- 
LOTYPE: PH presumably, fragment MO). 


Cuscuta eda Behr, Proc. Cali i ae (ed. 2) 1:16. 1854. The protologue he no ormariOn 


anda probably will be required to firmly establish th 

tit = of this name (see comments ca) Cuscuta ceanothi is the earlier name but is tenta- 

tively placed here with C. subinclusa. 
Stems medium, sometimes fleshy, creamy. Inflorescences few- to several-[low- 
ered, in scattered to densely aggregated clusters, pedicels 0-1 mm long; bracts 
1-0, ovate to lanceolate. Flowers 5-merous, 5-7(-9 mm) long, white. Papillae 
present in the corolla lobes; laticifers isolated or in rows obvious in the calyx, 
corolla and fruit. Calyx campanulate, ca. 1/2 as long as the corolla tube, divided 
3/5-2/3 the length, lobes broadly ovate to lanceolate, acute, sometimes cuspi- 
date, basally overlapping. Corolla tube cylindric, 2.5-3.5(-4.5) mm long, usu- 
ally showing horizontal ridges between the stamen attachments when dry, lobes 
ovate-triangular, acute and often slightly acuminate, 1/4-1/3 as long as the tube, 
widely spreading to reflexed. Stamens subincluded; anthers linear 0.8-2 mm; 
filaments 0-O.1 mim. Pollen 3(-4)-zonocolpate (15-)17-22(-24) um long, poly- 
morph, subsphaerical to subprolate, rounded at poles, tectum microreticulate, 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 187 


Fic. 3. Morphology of is.a. C. californica (var. californica), lateral view. b. C. occidentalis, lateral view. c. C howelliana 
lateral view (scale bar = 0.38 mm). d. Hilum of C. californica. e—g. C. howelliana: e. hilum, f-g. variation of seed coat 
micromorphology (scale bar = 100 um). 


188 BRIT.ORG/SIDA 22(1) 


puncta, 0.4-0.9 tum in diameter, granulate (Fig. 2 ef). Infrastaminal scales ob- 
long to spatulate, irregularly short-fimbriate, ca. 1/2 as long as the corolla tube; 
styles 1-1.5 mm long; stigmas capitate, globose. Capsules ovate to elliptical, 1.5- 
3 x 1.2-2.5mm, pointed, thickened in the form of a collar about the interstylar 
aperture, indehiscent or irregularly dehiscent, capped by the withered corolla. 
Seeds mostly | per capsule, not visible through the capsule, subglobose to 
broadly ovoid, rarely slightly dorsoventrally compressed, 1.3-1.7 x 1.2-1.5 mm; 
hilum terminal; seed coat cells alveolate/papillate when dried and papillate 
when hydrated, cells 30-40 um in diameter. n = 14 (Beliz 1986). 

Distribution and ecology—U.S.A.: California, Oregon. MEXICO: Baja Cali- 
fornia. Flowering Apr-Oct. Hosts: wide variety of woody and sometimes her- 
baceous plants growing along river banks and canyon bottoms, sometimes in 
salt marshes and deserts, including species of Adenostoma, Amelanchier, Arc- 
tostaphylos, Artemisia, Asclepias, Ceanothus, Cercis, Citrus, Clematis, Erigeron, 
Grindelia, Heteromeles, Monardella, Rhus, Schinus, Populus, Rhododendron, 
Rosa, Salix, Solidago, Vitis, particularly common on Rhus and Eriogonum. 

Conservation status.—G4 (apparently secure) (not yet assessed by 
NatureServe 2005). 


4. Cuscuta suksdorfii Yuncker, Mem. Torrey Bot. Club 18:167. 1932. Type: US.A 
/ ASHINGTON. Skamania Co. on an island of a mountain lake, on Aster, 24 Sep 1891, Suksdorf 

1487 (HOLOTYPE: US, fragment NY, IsoTypEs: GH, NY-2 sheets). This was essentially a renam- 

ing at varietal rank of Cuscuta salina var. acuminata (see below), but apparently Yunc 


— 


Ker 
intended for it to be taken as a homotypic new species. 

Stems slender, yellow. Inflorescences few-flowered umbellate clusters: pedicels 
0-2 mm long; bracts 0-1, ovate lanceolate. Flowers 4-5-merous, white, 2.8-3.3 
mm long; papillae absent; laticifers isolated in the calyx, corolla and ovary/ 
capsule. Calyx broadly campanulate often zygomorphic, reaching ca 1/2 tothe 
corolla lobes tips, divided 1/2-3/5, lobes ovate more or less unequal, with long- 
attenuate tips, not basally overlapping (Fig. le). Corolla 2.9-3.2 mm long: tube 
campanulate, 1.2-1.5 mm, lobes triangular-ovate, with lanceolate-attenuate tips, 
longer than the tube, suberect. Stamens included or barely visible through the 
corolla sinuses, anthers broadly elliptic, 0.2-0.4 mm long, filaments longer than 
anthers. Pollen 3(-4)-zonocolpate 18-22(-24) ttm, polymorph, subsphaerical to 
subprolate, rounded at poles, tectum perforatum, puncta, 0.3-0.5 1m in diam- 
eter, granulate. Infrastaminal scales oblong, represented by shallowly toothed 
wings, |/2-3/4 as longas the corolla tube; styles distinct, terete to slightly subu- 
late, 0.3-0.7 mm long; stigmas capitate, globose. Capsules elliptical-ovoid, ovoid- 
conic, globose to depressed globose, 2-3.2 x 2-3.6 mm, irregularly dehiscent, 
with withered corolla surrounding lower half. Seeds 2-4 per capsule, 0.80-1.1 x 
0.8-1.02 mm, dorsoventrally compressed, subround, hilum subterminal, 0.25- 
0.3 x 0.2-0.28 mm, vascular scar, 0.04-0.07 mm, oblique; alveolate when dry 
and papillose when hydrated, cells 30-50 um in diameter. n = 14 (Beliz 1986). 


— 


—_ 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 189 


REY: 1O VARIETIES OF CUSCUIA SURSDOREL 


. Flowers on pedicels 0.5-2 mm; calyx 2.2-2.6 mm long; capsule ae to ovoid- 
conic 4a. C. suksdorfii var. suksdorfii 
. Flowers sessile or subsessile; calyx 1.6-2 mm long; capsule me to depressed 
glo 


eo 


4b. C. suksdorfii var. Serenata 


4a. Cuscuta suksdorfii var. suksdorfii. Cuscuta salina var. acuminata Yuncker, Illinois Biol. 
Monogr. 6, pt. 2, 3:72, fig. 32,89. 1921 (non C. acuminata Nutt. ex Engelm. 1859, in adnot.). TYPE: 
U.S.A. WASHINGTON. Skamania Co: on an island of a mountain lake, 24 Sep 1891, Suksdorf 
1487 (HOLOTYPE: US, fragment NY; IsOTYPES: CAS, GH, MO, NY-2 sheets). 


Distribution and ecology.—U.S.A.: California, Oregon, Washington. Flowering 
Jul-Sep. Hosts: herbaceous species, mostly Asteraceae, Calyptridium, Trifolium. 
Mountain meadows, 5000-8500 ft 

Conservation status.—G2G3 (imperiled to vulnerable) (t] 
in NatureServe 2005). 


4b. Cuscuta suksdorfii var. and Aare C uscuta suksdorfii var. sulpedieallcia Yuncker, 
Bull. Torrey Bot. Club 62:512. 1935. TyPE: U.S.A. CALIFORNIA. Siskiyou Co. Siskiyou Mts., Head E 
Fork Horse Creek, 6500 ft, 59 T47 N, R1O W, a Calyptridium umbellatum, 21 Aug 1934, 

L.C. Wheeler 3192 (HOLOTYPE: NY). 


Distribution and ecology.—U.S.A.: California, Flowering Jul-Sep. Hosts: 
Calyptridium. Mountain meadows, 6500 ft. 

Conservation status.—G2G3 (imperiled to vulnerable) (the same assessment 
in NatureServe 2005). 


5. Cuscuta californica Hook. & Arn., Bot. Beechey Voy., 364. 1839 (non Choisy 
1842). Type: Presumably in K, not designated by Hooker and Arnott. Engelmann (1859) com- 
mented that “Both [Choisy and Hooker & Arnott] described this plant from Douglas's speci- 

mens under the same name and in the same year ( )” De Candolle (Prodr. vol. 9, 1845, p. 
jes cited “hab. Nov. Californiam ubi rep. Douglas! h. soc. hort. Lond. Hook. bot. Beech. suppl 
ie 364.” Yuncker (1921, 1932) apparently repeated de Candolle’s citation, noting for the type 

v. Californiam’ (Douglas). Not seen.’ 


Stems —— yellow to orange. Inflorescences loose, cymose-paniculate clus- 
ters; pedicels (0.5)1-2.5(-3 mm); bracts lanceolate to ovate. Flowers 5-merous, 
white, 3-5(-6.5) mm long; papillae sometimes present on the pedicels, receptacle, 
calyx and corolla; laticifers isolated, elongated present in the corolla and capsules. 
Calyx turbinate-campanulate, sometimes together with the receptacle, fleshy at 
the base, 1/2 to ca. as long as the corolla tube, divided 1/2-1/3 to the base, lobes 
triangular to lanceolate, acute to acuminate, basally overlapping. Corolla persis- 
tent; tube campanulate-cylindric, 1.9-2.7 mm, sometimes showing horizontal 
ridges between the stamen attachments when dry, rarely somewhat saccate, lobes 
narrowly lanceolate, acute, shorter than, equaling or longer than the tube, ini- 
tially erect, later reflexed (Fig. 1a, b,c). Stamens + exerted when flowers are com- 
pletely open, anthers oblong to linear, 0.7-1.1 mm long, filaments 0.1-1.1 mm long. 
Pollen 3(-4)-zonocolpate (16-)17.5-22.6(-26) um, polymorph, sphaerical to pro- 


190 BRIT.ORG/SIDA 22(1) 


late (the latter more common), rounded at poles, tectum imperforatum or witha 
few puncta, 0.2-0.3 um in diameter, granulate. Infrastaminal scales completely 
lacking or reduced, 0.1 mm long at the base of the corolla tube. Styles distinct, 
evenly filiform, 12-3 mm long, stigmas capitate, globose. Capsules globose or 
ovoid-conic, 1.6-2.3 x 1.8-2.5 mm, sometimes apically pointed, indehiscent or ir- 
regularly dehiscent, completely hidden by the withered corolla (Fig. 1g); pericarp 
and persistent corolla not semi-transparent (thicker than in C occidentalis). Seeds 
1-4 per capsule, not visible through the persistent corolla and pericarp, 0.90-1.4 
x 0.85-1.2 mm, dorsoventrally compressed, broadly-elliptic to obovate, hilum 
lateral, inconspicuous, subround 0.10-0.15 mm, vascular scar 0.05-0,06 mm, ver- 
tical; entirely alveolate (hilum area included) when dry and papillose when hy- 
drated, cells 25-50 um in diameter (Fig. 3a, d).n = 14 (Beliz 1986). 

In some plants of Cuscuta californica (particularly of vars. apodanthera 
and apiculata), the receptacle and perianth cells located at the base of capsules 
and seeds are fleshy and become mucilaginous when brought in contact with 
water. A similar phenomenon occurs in C. nevadensis LM. Johnston and to a 
lesser extent in C. veatchii Brandegee; Costea et al. 2005 suggest these features 
may be related to vivipary observed in C. nevadensis. The fleshy receptacle and 
perianth base in C. californica may be homologous with those of C. nevadensis, 
but in contrast to C. nevadensis, C. californica does not have seeds with a globu- 
lar-enlarged embryo at the hilum end. Apart from scattered observations such 
as these, the reproductive biology of Cuscuta species is poorly known (reviewed 
by Dawson et al. 1994; Costea & Tardif 2006). 


KEY TO VARIETIES OF CUSCUTA CALIFORNICA 


. Ovary and fruit conic, apically pointed; 1-seeded 5b. C. californica var. apiculata 
eee and fruit globose to globose depressed; (1-)2-4 seeded. 
2. Flowers and pedicels densely papillate___ 5c. C. californica var. papillosa 
2. Flowers and pedicels not papillate. 

3. Anthers sessile or subsessile 5d. C. californica var. apodanther 

3. Anthers not sessile, filaments shorter to equaling anthers 5a.C. Seni 


var. californica 


5a. Cuscuta californica var. californica. Cuscuta californica Hook. & Arn. var. graciliflora 
Engelm., Trans. Acad. Sci. St. Louis 1499. 1859, LECTOTYPE (designated noe : oe CALIFOR- 
NIA: “Almost without flowers, on Monarda,” 


datal, 1846 (MO). Engelmann cited “California, Dotigiast Fremont! 506: ae 


Cuscutad ca omnes Hook. & Arn. var. li loba Engelm., Trans. Acad. Sci. St. Louis 1:499. 1859. 
LECTOTYPE (designated here): U.S.A. CALIFORNIA: San Felipe, on Eriogonum polifolium, Jan 1852, 
Thurber 633 (MO). Engelmann pee ‘Sta. Barbara, Nuttall! San Diego, Thurber! 570 & 633; 
Newberry!” Engelmann’s material (at MO) of the Nuttall collection apparently is a fragment 
of the son ee specimen at PH, judging from notes inside the packet 


Cus tlifornica Hook. & Arn, var. brachycalyx Yuncker, ite Biol. Monogr. 6, nos. 2, 3:62, 


fig. 45, 75, 51921 Type: U'S.A. CALIFORNIA: Near Hanford, dry soil on Centromadia pungens, 21 
Jun L901, Kearney 52 (HOLOTYPE: NY; ISOTYPE: US). 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 191 


Cuscuta brachycalyx Yuncker, Mem. Torrey Bot. Club 18:159. 1932. Type: U.S.A. Cam IFORNIA: Near 
Hanford, dry soil on Centromadia pungens, 21 Jun 1901, Kearney 52 ( ). Yuncker 


treated this as a new species, even though he cited C. californica var. br ee in synonymy. 


Distribution and ecology.—U.S.A.: Arizona, California, Nevada, Oregon, Utah, 
Washington; Mexico (Baja California). Flowering Mar-Aug(-Sep). Hosts: 
Abronia, Adenostoma, Agastache, Ambrosia, Asclepias, Convolvulus, Corethrogyne, 
Croton, Eriodictyon, Eriogonum, Holocarpha, Iva, Lupinus, Salvia, and others. 
200-22650 feet. 

Conservation status.—T5 (common) (the same assessment in NatureServe 
2005) 


5b. Cuscuta californica var. apiculata Engelm., Trans. Acad. Sci. St. Louis 1:499. 
TYPE: U.S.A. CALIFORNIA: “Dry arroyos, on Dalea spinosa, [on the] Colorado,” 22 Feb 1884, 
J.M. Bigelow s.n. (HOLOTYPE: MO; ISOTYPES: GH, NY). 
Distribution and ecology—U.S.A.: California, Nevada. MEXICO: Baja California. 
Flowering Mar-Aug. Hosts: various species in sandy desert areas. 
Conservation status.—T1T2 (critically imperiled to imperiled), (T3?, 
Natureserve 2005). 


5c. Cuscuta californica var. papillosa Yuncker, Illinois Biol. Monogr. 6:152. 1921. 
TYPE: U.S.A. CALIFORNIA. San Bernardino Co. San Bernardino Valley, Parish 5524 (HOLOTYPE: 


el 


Distribution and ecology.—U.S.A.: California (endemic): Imperial, Riverside, San 
Bernadino, and San Diego cos. Flowering Mar-Aug, -Sep. Hosts: various species 
in sandy desert areas. 300-4600 feet. 

Conservation status.—T3 (vulnerable) (the same assessment in NatureServe 
2005) 


5d. Cuscuta californica var. apodanthera Yuncker, Illinois Biol. Monogr. 6:152. 
1921. Cuscuta brachycalyx var. apodanthera (Yuncker) Yuncker, Mem. Torrey Bot. Club 18:159. 
1932. Type: U.S.A. CALIFORNIA: Yosemite Valley, 7-12 Jul 1896, Jepson 80a (HOLOTYPE: JEPS, frag- 
ment NY). 


Distribution and ecology.—U.S.A.: Endemic in California. Flowering Aug-Sep. 
Hosts: Eriogonum and other herbs. 

Conservation status.—T1T2 (critically imperiled to imperiled) (T2, 
NatureServe 2005). 


6. Cuscuta occidentalis Millsp. nom. nov. Bee Field Columbian Mus., Bot. Ser. 
204.1923. Based on C Arn. var. breviflora Engelm., Trans. Acad. 
Sci. St. Louis 1:499. 1859. Grammnicd Bei (Millsp.) cca & Chrtek, Folia Geobot. 
Phytotax. 8:220. (Aug) 1973. Grammica occidentalis (Millsp.) W.A. Weber, Southw. Naturalist 
18:319. (Oct) 1973. TYPE: U.S.A. CALIFORNIA. Monterey Co.: rae in fields, [no date], Hartweg 

1863 (HOLOTYPE: MO; ISOTYPES: GH, NY) 


Stems medium, yellowish to orange. Inflorescences compact glomerulate clus- 
ters; pedicels 0-0.5(-1.5 mm); bracts lanceolate to ovate. Flowers 5-merous, white 


192 BRIT.ORG/SIDA 22(1) 


to creamy; 2.7-3.4 mm long; papillae rarely present; laticifers not conspicuous 
(Fig. 1d). Calyx campanulate, somewhat shorter to ca. as long as the corolla tube, 
divided 2/5-1/2 the length, rarely fleshy at base, lobes narrowly ovate to lan- 
ceolate, acuminate, not basally overlapping; corolla tube cylindric-campanu- 
late, 1.4-2.1 mm long, saccate between the stamen attachments, lobes lanceolate, 
acuminate, shorter than the tube, initially erect, later usually spreading (some- 
times reflexed) giving flowers a star-shaped appearance. Stamens + exserted; 
anthers broadly-elliptic, 0.25-0.5 mm long, filaments 0.3-1 mm long. Pollen as 
in C. californica, 17-)19-24(-26) um long. Infrastaminal scales as in C. 
californica; styles distinct, evenly filiform, styles 0.5-1(-L5) mm long; stigmas 
capitate, globose. Capsules globose to globose-depressed, 1.8-2.2 x 2-2.6 mm, 
indehiscent or irregularly dehiscent, not completely enclosed by the corolla (the 
top of capsule visible) (Fig. 1h); corolla and pericarp thin, membranous and 
semi-transparent. Seeds 2-4 per capsule, visible through both corolla and peri- 
carp, 0.85-L3 « 0.8-1.1, dorsoventrally compressed, subround to broadly-ellip- 
tic, hilum subterminal, poorly differentiated from the rest of the seed, subround 
0.10-0.15 mm, vascular scar 0.05-0.06 mm, vertical, surface of seed coat alveo- 
ate when dry (including hilum area), cells 25-50 um in diameter (Fig. 3b). 

Distribution and ecology.—U.S.A.: California, Colorado, Idaho, Nevada, Or- 
egon, Utah, Washington, Wyoming. MEXICO: Baja California. Flowering Mar- 
Aug(-Sep). Hosts: Artemisia, Boisduvalia, Calyptridium, Corethrogyne, 
Ericameria, Eriogonum, Hemizonia, lva, Lotus, Lupinus, Monardella, Oenothera, 
Polygonum, Salvia, Sisymbrium, and others. 25-6500 feet. 

Conservation status.—G4G5 (apparently secure to secure) (not assessed yet 

by NatureServe 2005). 


APPENDIX 1.~VOUCHERS FOR THE SEM STUDY 
Vouchers are from NY unless otherwise indicated. 
1. Cuscuta salina (see Costea et al. 2005). 
2.Cuscuta howelliana (7 collection d).— U.S.A. CALIFORNIA. Butte Co.: ca 7 mi N 
of Oroville, Table Mountain, E side of Cherokee Rd., 1325 ft, 18 Jul 1998, Ahart 6044 (JEPS). Sacra- 
mento Co.: ca 1 mi S of White Rock Rd. and Scott Rd. intersection, 27 May 1985, Bowcutt 337 (UC). 
Shasta Co.: between Goose Valley and Burney Valley, ca. 3.5 mi NNW from Burney, 3200 ft, 8 Aug 
1988, Taylor 10026 (JEPS). Siskiyou Co.: 16 mi N of Fall River Mills (Shasta Co.) and 6 mi W of Day 
(Modoc Co.), 1.2 mi E of Spring Creek Rd., 3600 ft, 19 Jul 1989, Powell 3533 EPS). Tehama Co.:ca 10 
mi SE of Corning, S Fork Hall Creek, 460 ft, 12 May 1995, Taylor 14897 (JJEPS); BLM parcel along Hwy 36 
NE of Red Bluff, 870 ft, 8 Jul 1996, Oswald & Ahart 7978 (JEPS): Inks Creek, ca. 1.5 mi N of Dales Lake, 
612 ft, 27 May 1992, Taylor 12661 (UC). 
3. Cuscuta subinclusa (11 collections examined). oe S.A. CALIFORNIA. [no county given, 
Los Angeles or San Bernardino Co.?] Swartout, 1 Sep 1939, Grace s.n. Los Angeles Co.: Liebre 
Mts., entrance to Ruby Canyon of Forest ae Rd., ca. : 0 mi from junction with Lake Hughes 
Rd., 2000 ft, 23 Oct 1996, Raz & Boyd 015; Lower Spunky Canyon, stretch draining SSE exiting into 
Bouquet Reservoir, 2995-3065 ft, 20 Sep 1994, Ross & Boyd 8260; San Gabriel Mts., Tujunga Creek, 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 193 


near mouth of Canyon, 23 Aug 1937, Ewan 11049. Marin Co.: mouth of San Antonio Creek, 3 Sep 
1939, Howell 15355; Mouth of San Gabriel Canyon, 17 Dec 1935, Hastings s.n.;San Antonio eo N 
of Novato, 5 ft, 3 Sep 1939, Rose 39363. eOner ey Co.: Banks of San Antonio River, Santa Lucia Mts., 
25 Oct 1930, Mason 5766.Nevada Co.: Banner Mt.,ca.4 mi E of Nevada City, 3900 ft, 15 Jul er True 
& Howell 2315. Plumas Co.: Mill erealiun 1877, Austin s.n. San Joaquin Co.: San Joaquin Bridge 
near Lathrop, 9 Sep 1892, Brandegee s.n 
4.Cuscuta suksdorfii (5 collections examined).—U.S.A. CALIFORNIA. Humboldt Co:: Trin- 
ity Summit, near Box Camp, 5000 ft, 4 Aug 1949, Tracy 18430 (UC). Siskiyou Co.: Siskiyou Mts., M 
Diablo, 1 mi E Dry Lake Lookout, 6000 ft, 31 Jul 1934, Wheeler 3011 UJEPS);.[Co. inepecieal ae 
Canyon National Park, Woods Creek, near base of Castle Dome, 8500 ft, 27 Jul 1948, Bailey & Bailey 
2672a (UC). [Co. unspecified] ca 1/4 mi E of Yuba Gap and Hwy 80, E side of the paved road to Lake 
Valley Reservoir, 5840 m, 22 Jul 2002, Ahart 9885 (JEPS). [No location, no date, no collector] (UC 
23290). 
5. Cuscuta uae 
5a. uta californica var. californica (13 collections examined).—U.S.A. CALIFORNIA. 
[locality nee 21 Jun 1931, Yuncker & Welch 3748. Kern Co.: “brachycalyx,” 1.5 mi N of Kernville, 
ep 1962, Howell 38877. Las Angeles Co.:Liebre Mts., Knapp Ranch at the ee end of 
Castaic Creek drainage in pio alluvial valley at head of Cienaga Canyon, S of Liebre 2840- 
3080 ft, 28 May 1997, Boyd 9839; [some flowers “brachycalyx"-like] Whittier Hills, S- aan see 
of Turnbull Canyon, 980-1000 ft, 15 Jun 1992, Ross 6575; Mount Wilson, 3000 ft, 17 Jul 1931, Yuncker 
& Welch 3704, 3707. Orange Co.: Santa Ana Mts., Silverado Canyon, 21 Jun 1931, Fosberg 5143. River- 
side Co.: 8 mi SE of Corona, along road to Elsinore, 17 Mar 1964, Hitchcock & Muhlick 23076. a 
Bernardino a foot hills of San Bernardino Mountains, 15 Jun 1898, Parish s.n. San Diego Co. 
Jamul Valley, Jun 1875, Palmer 439; San Diego, Mission Hills, 14(?) May 1903, Abrams 3462; along re 
78E 5.3 Mi W of Ramona, 15 Jun 1973, LeDoux et al. 114; [some flowers “brachycalyx”-like] Coronado 
Beach, 24 Jun 1932, Moldenke 7037. San Luis Obispo Co.: Pismo, 25 Aug 1932, Demaree 9281.NE- 
VADA. Clark Co.: Hells Kitchen area near Gold Butte, Devils Cove Rd., 2.9 mi S of Connoly Spring, 
2400 ft, 14 May 1986, Pinal 72 
5b. Cuscuta californica var. iepapillosa (5 collections examined).—U.S.A. CALIFORNIA. [no 
a Aug 1901, Grant 4449. Riverside Co.: Cathedral City, 23 Apr 1945, Lillian s.n.; ca. 500 ft, 27 
1945, Rose 45331; Cathedral Canyon, 23 Dec 1946; Rose 46300. San Bernardino Co.: 1/2 a E of 
ae Ranch, Horsethief Canyon, Mohave River tributary, 3300 ft, 17 Sep 1933, Wheeler 271 
Cuscuta californica. var. apiculata (3 collections examined).—U.S.A. PAIEORAIA Riv- 
erside Co.: Cathedral City,no date (probably 1945), Blake s.n. NEVADA. Clarke Co.: 4 mi SE of Muddy 
Peak at head of Callville Wash, 2500 ft, 28 May 1937, LaRivers & Hancock 261; 29 May 1937,LaRivers & 
Hancock 231 


ta identalis (14 collections examined).—U.S.A. Tee Fresno Co.: Cop- 

per Creek Canyon Creek Trail, 7200 ft, 2 Aug 1958, Howell 34198.Mono Co.: 1 mi Eof Mammoth Post 
Office, 29 Jul 1951, Raven 3720.Monterey Co.: Point Lobos State Park, 16 Jun ees. 9219. 
Siskiyou Co.: along Klamath River Hwy.W of Pacific Hwy., 2000 ft, 20 Jun 1940, Yuncker 15144. Trin- 
ity Co.: Cold Springs Area, South Fork Mt.,5400 ft, 28 Aug 1941, Parks & Tracy 11517. Washoe Co. 
1 mi ENE of Grass Valley Reservoir, 6000 ft, 14 Aug 1991, Schoolcraft et al. 2220. COLORADO. Duis 
oe Paonia, 23 Jul 1911, Osterhout 4602.IDAHO. Adams Co.: 2 Mi S of Council, 26 Aug 1937, Christ & 

ard 8787.Gooding Co.: Little City of the Rocks in the Mt. Bennett Hills N of Gooding, 28 Jul 1976, 
eae NEVADA. Douglas Co.: Antelope Valley at the E edge of Red Hill on the E side of Gray 
Hills, 4990 ft, 14 Jul 1997, Tiehm 12257. Eureka Co.: Roberts Mts., 1.7 mi N of Three Bars Ranch on the 
main road to Tonkin Summit, 6600 ft, 22 Aug 2002, Tiehm 14108. Mineral Co.: E slope of Wassuk 
Range, base of Big Indian Mt., 9500 ft, 11 Sep 1938, Archer 7032. Washoe Co.: dry bed of Ice Pond, 
Whittell-Audubon Area, Carson Range foothills, 5480 ft, 12 Aug 1974, Howell et al. 50745.UTAH. Salt 
Lake Co.: [no locality], 16 Aug 1907, Garrett 2170. 


194 BRIT.ORG/SIDA 22(1) 


ACKNOWLEDGMENTS 


We thank directors and/or curators from ACAD, ALTA, ARIZ, ASU, BRIT, DAO, 
F GH, HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, 
Roa opoK ols, (EA Oo LE.TUP UBC, UC EPS. UNE, UNM, US USis.U WO: 
UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Special thanks are 
due to NY staff for approving and preparing the four voluminous loans con- 
taining the herbarium of T.G. Yuncker. Dan Austin and an: mous reviewer 
provided useful comments and suggestions for an earlier version of the manu- 
script. Thierry Deroin kindly donated all the Cuscuta fragments annotated by 
Yuncker (held in P) to WLU. Barbara Ertter (UC) sent us copies of the relevant 
publications of Behr and Currant. Alexandra Smith assisted us with the scan- 
ning electron microscope and Gabriel Moreno-Hagelsieb translated the Span- 
ish abstract. 


REFERENCES 


BeHr, H. 1888. Flora of the vicinity of San Francisco. San Francisco, California (privately 
published). 

Beuz, T. 1986.A revision of Cuscuta sect. Cleistogrammica using phenetic and cladistic analy- 
ses with a comparison of reproductive mechanisms and host preferences in species 
from California, Mexico, and Central America. Ph.D. diss., Univ. of California, Berkeley. 

Betiz, T. 1993. Cuscuta.|n: J. Hickman, ed. Jepson’s manual: higher plants of California. Univ. 
of California Press, Berkeley. Pp. 538—539. 

Costea, M. and FJ. Taroir. 2006. Biology of Canadian weeds. Cuscuta campestris Yuncker, C. 
gronovii Willd. ex Schult., C. umbrosa Beyr.ex Hook., C. epithymum (L.) L.and C.epilinum 
Weihe. Can. J. Pl. Sci. 86:293-316 

Costea, M.,G.L.Nesom, and FE. Tarpir.2005. Taxonomic status of Cuscuta nevadensis and Cuscuta 
veatchii (Convolvulaceae). Brittonia 57:264-272. 

Costea, M., G.L. Nesom, and S. Steranovic. 2006a. Taxonomy of the Cuscuta pentagona com- 
plex (subsect. Arvenses: Convolvulaceae) in North America. Sida 22:151-175. 

Costea, M., G.L. Nesom, and S. Steranovic. 2006b. Taxonomy of Cuscuta gronovii and Cuscuta 
umbrosa (Convolvulaceae). Sida 22:197-207. 

Costea, M., G.L. Nesom, and S. Steranovic. 2006c. Taxonomy of the Cuscuta indecora 
(Convolvulaceae) complex in North America. Sida 22:209-225. 

Curran, M.K. 1885. List of the plants described in California, principally in the Proc. of the 
Calif. Acad. of Sciences, by Dr. Albert Kellogg, Dr. H.H. Behr, and Mr. H.N. Bolander: with 
an attempt at their identification. Bull. Calif. Acad. Sci. 1:128-151. 

Dawson, J.H., LJ. Musseuman, P.WotswinkeL, and |. Dorr. 1994. Biology and control of Cuscuta. 
Rev.Weed Sci. 6:265-317. 

ENGELMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with 
critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 
1:453-523. 


COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 195 


ENceLMANN, G. 1876. Cuscuta salina. In: W.H. Brewer, S. Watson, and A. Gray. The botany of 
California, Univ. Press, Cambridge. P. 536 

Ertter, B. 2003. Mein lieber herr College: letters from Behr to Engelmann. Fremontia 31: 
18-22. 

NatureServe. 2005. NatureServe Explorer:An online Sneyclopediar of life [web pplication! 
Version 4.0, NatureServe, Arlington, Virginia. <http:// ve.org/exp = 
(Accessed: November 16, 2005). 

Pazy, B. and U. Putmann. 1995.Chromosome divergence in the genus Cuscuta and its sys- 
tematic implications. Caryologia 48:1 73-180. 

Ruetzorr, P 1966.A new Cuscuta from California. Leafl. West. Bot. 10:335-337. 

Yuncker, T.G. 1921 (1970). Revision of the North American and West Indian species of 
Cuscuta. Illinois Biol. Monogr. 6:91-231. 

YUNCKER, T.G. 1932. The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. 

YUNCKER, T.G. 1965. Cuscuta. North American flora, ser. 2,4:1-51. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Fa-Ti Fan. 2004. British Naturalists in Qing China: Science, Empire and Cul- 
tural Encounter. (ISBN 0-674-01143-0, hbk.) Harvard University Press, 79 
Garden St. Cambridge, MA. (Orders: http://www.hup.harvard.edu/; 800- 
405-1619; email: Contact|HUP@harvard.edu). $52.00, 238 pp, Lo illustra- 
tions, 91/2" X 61/4". 

The Qing (Manchu) Dynasty lasted nearly 300 years oo ru) if me West it was He period of the 

Industrial Revolution, the rapid growth of scientific | 

alism of the 17" and 18" centuries. China had had its own economic revolution between the Bi and 

12" centuries, but the potential of the great advances made at that time in agriculture, techno ology, 


large-scale industry, and the use ol money and credit was reaching its end. So the two cultures were 
at different stages in the Confu the rise and fall of states when they met. 


Although the Chinese reaction 6 the increasing Western presence was to restrict foreign trade 
toa single port an enormous volume of Chinese goods Bene flora and fauna) was ap Uno to 
‘ oO 


Europe. It not only affected European taste but 
This book, written from the viewpoint of a modern historian, traces the activities of the British natu- 
ralists within the cultural context, examining the institutional base of enterprise as well as the pur- 
poses, methods and results for each side. Because the ee became predominant in the China trade, 
they were best able et the Chinese was essential to their work. The author 
insists that it is the ae hybridization, and confluence’ acrossa variety of borders and barriers 
that made possible the great advances in ethno-botany, economic botany and Sinology of this pe- 
ue _— e es oS not downplay the role of British HETWOES of MESS dipl omats, missionaries, 
I | previously unacknowledged) 


a 


eographers, and geologists 
em i . to the part eed by the indigenous people the cen merchants, compradors, ser- 
vants, gardeners, craftsmen, artists, etc. His aim is to “explain the formation of scientific practice and 


knowledge in cultural borderlands during this critical period.” 

The book naturally divides itself into the time before and ans op ee War. Peeve ss 
abrupt policy changes which followed granted W g g I 
increased access to the interior of the country, there I in the procedures. Both 
sections are extremely interesting and well- researched. An excellent fades copious notes, short bi- 
ographies of the many British naturalists mentioned, maps and illustrations all add to the value of 
the work. British Naturalists in Qing China offers fresh insights into the very many aspects of Sino- 
British relations. It is particularly timely as China emerges as a world power—Ruth Ginsburg, Botani- 
cal Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. 


— 


SIDA 22(1):196 196. 2006 


TAXONOMY OF CUSCUTA GRONOVII AND 
CUSCUTA UMBROSA (CONVOLVULACEAE) 


Mihai Costea (corresponding author) Guy L.Nesom 
Department of Biology Botanical Research Institute of Texas 
Wilfrid Laurier University 509 Pecan Street 
75 University Avenue West Fort Worth, Texas 76102-4060, U.S.A. 
Waterloo, Ontario gnesom@brit.org 


N2L 3C5, CANADA 
mecostea@wlu.ca 


Sasa Stefanovic 


Department of Biolo 
University of Toronto at Mississauga 
Mississauga Road 
Mississauga, Ontario 


L5L 1C6, CANADA 


ABSTRACT 


Cuscuta Pies is the correct name for the species often recognized as C. megalocarpa. It appar- 
ently i e features with C. gronovii, but the two are generally distinct across a broad 
region of ee ies varieties are recognized within C. gronovii: var. gronovii is widespread; var. 
latiflora is broadly sympatric with var. gronovii but is maintained because relationships with C. 
gronovii var. gronovii and C. cephalanthi are not clear; var. calyptrata is more distinct and narrowly 
restricted in range, although it also is sympatric with var. gronovii. 


RESUMEN 


Cr 
E li i Atri lgoi g dad eronovii lasd 


P Seeecat =) 
teandadec Apntrnde 


Cc Fs ] 1 | ] ; ore i C. meealocarpa 
ee ym Bs 
7 ] t 


o Q y o 
gronovii: var. gronovii que esta muy esparcida; var. 
latiflora es ampliamente simpatrica con la var. gronovii pero se mantiene debido a la relacion con C. 
& 
gronovii var. Ses y C.cephalanthi no esta clara; la var. calyptrata es mas clara y estrechamente 


restringida ribuciér geografica aunque sea también simpatrica con la var. gronovit. 


L A 
claras. Se reconocen tres 


The original concept of Cuscuta umbrosa Beyr. ex Hook. (1840) included C. 
gronovii Willd. ex Roem. & Schult. 1820) in typical form as well as a variant 
plant, based on the specimens cited in the protologue. Engelmann (1859) clari- 
fied the typification of C. umbrosa, distinguishing it from C. gronovii, and de- 
scribed the variant as C. gronovii var. curta Engelm. Yuncker recognized the 
variant taxon at specific rank, at first (1921, 1932) using the name C. curta 
(Engelm.) Rydb. but later (1943, 1965) adopting C. umbrosa as the earliest legiti- 
mate name. We find that recently C. umbrosa either has been accepted as a spe- 
cies under yet another name, C. megalocarpa Rydb. (1901) (Kartesz 1999; 


SIDA 22(1): 197 — 207. 2006 


198 BRIT.ORG/SIDA 22(1 


= 


Darbyshire 2003; USDA, NRCS 2004), or else it has not been recognized at any 
taxonomic rank (Beliz 1986-1987, in herb; Crins & Ford 1988). 

Cuscuta gronovii and C. umbrosa, as recognized here, are similar and ap- 
parently closely related to C. cephalanthi Engelm. and C. rostrata Shuttlw. ex 
Engen & A. Gray. These four species, which were placed by Yuncker (1965) in 
two subsections (Platycarpae and Cephalanthae), have the same pattern of stem 
growth and branching: haustoria are formed as the main stem twines around 
the host and no tendril-like axillary branches are generated. This pattern was 
first observed by Dawson (1984) in C. gronovii. The evolutionary and taxonomic 
status of C. umbrosa is reevaluated here, especially with regard to its relation- 
ship with C. gronovii. 


METHODS 


Descriptions of morphology are based on samples from specimens from DAO, 
NY, and SMU (Appendix 1). Measurements and pictures were taken witha scan- 
ning electron microscope Hitachi S-570 at 15 KV. Samples were coated with 30 
nm gold using an Emitech K 550 sputter coater. Terminology regarding the mi- 
cromorphology of flowers, seeds and capsules, and pollen were described in 
detail in the first paper published in this issue (Costea et al. 2006). Conserva- 
tion status was assessed using NatureServe (2005) ranks and criteria. 


TAXONOMY 


Yuncker (1932, 1965) characterized Cuscuta gronovii as having “styles about 
equaling the ovary or somewhat shorter,” in contrast to those of C. umbrosa, 
which are “very short, about one fourth the length of the ovary.” Other con- 
trasts between these taxa emphasized by Yuncker are in the morphology of 
infrastaminal scales and the size of capsules and seeds. According to him, C. 
umbrosa has infrastaminal scales about half the length of the corolla tube, broad, 

often truncate and bifid, and capsules and seeds are larger. In contrast, C. gronovii 
has infrastaminal scales about equaling the corolla tube, oblong and not trun- 
cate or bifid, and capsules and seeds ar p ay smaller. We oe that C. 
gronovii and C. umbrosa usually are morphologically disting ; Dil G: 
gronovii may show character states similar to C. umbrosa even when it occurs 
outside the geographical range of the latter. For example, C. gronovii from On- 
tario and Québec may occasionally have more or less truncate infrastaminal 
scales one-third the length of the corolla tube, or short stigmas, or comparably 
large seeds (Crins & Ford 1988) and capsules. Commonly only one state char- 
acter of C. umbrosa, or rarely acombination of two (eg, large capsules and seeds), 
occurs in C. gronovii. Such plants of C. gronovii may be separated from C. umbrosa 
using the character combinations in the following key. Other differences can 
be found in pollen morphology: C. gronovii has pollen grains with tectum 


COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 199 


imperforatum or a few puncta, while C. umbrosa has tectum perforatum or al- 
most microreticulate (Fig. 1). 


. Infrastaminal scales oblon ly rounded, not bifid, rarely truncate, ca. reach- 
ing the filament bases, a ca. 1/3 Sp the Sole tube; styles (0.6-)1.2-2.2 mm 
long; capsules 2.5-4.5(-5.2) X 2-4(-5) mm; seeds 1.35-1.7(-2.4) * 1.2-1.5(-1.6) 


— 


C. gronovii 


1. 
. Infrastaminal scales broadly-oblong, apex truncate to slightly bilobed, 1/2(-1/3) as 
long as the corolla tube; styles 0.3-0.7(-0.9) mm long; capsules 3.5-6.5(-7) * 3-5 
(-6) mm; seeds (1.8-)2-2.5(-2.8) X 1.5-1.65 mm 2.C. umbrosa 


— 


Cuscuta gronovii has a wide geographic distribution in North America and is 
broadly sympatric with C. umbrosa. The former is more common in central and 
northeastern regions (Yuncker 1932), while the latter is primarily a prairie species. 
Cuscuta gronovii usually is distinct from C. umbrosa by a greater morphological 
gap than are the varieties of C. gronovii among themselves (see key below). 
Within Cuscuta gronovii, Yuncker (1965) recognized three varieties: var. 
gronovii, var. latiflora Engelm., and var. calyptrata Engelm. Variety latiflora and 
variety gronovii are broadly sympatric across their geographic range, they show 
no distinct preferences toward hosts, and morphological intermediates occur. 
Var. calyptrata is more narrowly restricted in range (Texas and Louisiana) but 
is sympatric with the widespread var. gronovii. Continued recognition of sym- 
patric varieties within Cuscuta gronovii is unusual. It reflects the observations 
that (1) a close morphological similarity exists among the varieties, (2) each of 
the varieties usually is morphologically distinct although intermediates may 
be relatively common, and (3) each of the non-typical varieties shows features 
suggesting that it might be more closely related to species of the Cuscuta 
cephalanthi-rostrata group or at least that genetic interchange among taxa of 
this broader group may have influenced the variation patterns within C. 
gronovii. We consider it p to discard the varieties as formally recognized 
taxonomic entities until relationships involving all these taxa are clarified. 


1. Cuscuta gronovii Willd.ex Roem. & Schult, Syst. Veg. 6:205. 1820 (Fig. 1 a, c, 
Epithymum gronovii (Willd. ex Roem. & Schult.) Nieuwl. & Lunell, Amer. Mid]. Naturalist 
4511. 1916. TyPE (see Yuncker 1943): North America, unspecified locality, Willdenow 3160-10 

(B, a photo seen) 


Stems medium to coarse, yellow to orange. Inflorescence of pedicellate (1-4.5 
mm) flowers in loose or dense, paniculate cymose clusters, sometimes endog- 
enously formed; each flower with one ovate to broadly triangular bract at the 
pedicel base, and sometimes 1-2(-4) bracteoles on the pedicels. Flowers 5-merous 
(sometimes 3- or 4-merous), 2-4(-4.7) mm long, not papillose, white-cream, 
with a few to many isolated, round to ale translucent laticifers in the calyx, 
corolla, and ovary. Calyx cupulate row-campanulate, reaching 
to ca. the middle of the earl aubed or the corolla sinuses, divided ca. 1/2-2/3 


| “ab I + +h L 


aronovil: b. CG 
g ! 


BRIT.ORG/SIDA 22(1) 


Fic. 1.a—b. C g 


1 c W “ 


ony la | 


7.5um 


pictures) (scale bar = 1 mm); c—e. Morphology of | 


g \ 
f. C gronovii: q. C. 
g g 


= 


/ 


+d-e, Cuscuta umbrosa 
1 1 nc \h 


WT), 


| 8 Sand 3m. resnectively):€—h 


Hydrated seed coat of C. umbrosa (scale bar = 120 Lim). 


COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 201 


the calyx length, lobes ovate to suborbicular or oblong, apex rounded or ob- 
tuse, more or less basally overlapping, margins entire to serrulate; corolla tube 
campanulate, broadly- or narrow campanulate (1-)1.5-2.5(-3) mm long, lobes 
mostly ovate, rounded-obtuse, commonly 1/3 the tube corolla tube, sometimes 
1/2 to equaling the corolla tube, spreading or reflexed; epicuticular wax absent 
or reduced to a few reticulated rodlets; stamens exserted, anthers on filaments 
equaling or longer than anthers; pollen 3(-4)-zonocolpate, (18-)19-24(-27) um 
long, rounded or truncated at poles, polymorphic in the same anther or flower, 
from spheroidal to subprolate (the former shape more frequent); tectum 
imperfortatum with a few puncta, rarely tectum perforatum with puncta 0.3- 
0.5 um in diameter; sexine scabrate usually with isolated granules; infrastaminal 
scales oblong or broadly-oblong, rounded or truncate and bifid, deeply fringed 
distally, shorter than the corolla tube or reaching the filament bases; styles dis- 
tinct, slender, thickened at the base or occasionally slightly subulate, (0.6-)1.2- 
2.2mm long; stigmas capitate, globose. Capsules globose, ovoid to globose-conic 
or subobpyriform, thickened at the top, interstylar opening relatively small, 
sometimes with a short and broad but discernible neck, 2.5-4.5(-5.2) x 2-4(-5) 

mm, indehiscent or irregularly dehiscent, surrounded or capped by the with- 

ered corolla. Seeds 2-4 per capsule, 1.35-1.7(-2.4) x 1.2-1.5(-L6), dorsoventrally 

compressed to obscurely angled, subrotund to broadly-ovate; hilum subtermi- 
nal, subrotund, 0.33-0.39 x 0.30-0.33 mm, vascular scar of funiculum linear, 
0.20-0.25 mm long, vertical to slightly oblique. Surface of seed epidermis vari- 
able: alveolate when dry and papillate when hydrated (cells 20-50 um in di- 
ameter), or cells more or less polygonal and puzzle-like, with epicuticular wax 
organized as reticulated rodlets. 2n = 60 (Fogelberg 1938). 


KEY TO VARIETIES OF CUSCUTA GRONOVIL 


— 


. Calyx tube about equaling corolla tube; calyx lobes oblong to ovate, barely over- 

lapping, corolla tube 1-1.5 mm long, broadly campanulate, corolla lobes ca. 1/2 

equaling corolla tube b. C. gronovii var. latiflora 
Calyx ca. 1/2 of corolla tube, calyx lobes ovate to suborbicular, overlapping; corolla 

tube 1.5-2.5 (-3) mm long, campanulate to narrow-campanulate; corolla lobes 1/ 

3(-1/4) of the corolla tube 

2. Corolla tube narrow-campanulate, 2-2.5 (-3) mm long, withered corolla cap- 


— 


ping capsule c. C. gronovii var. calyptrata 
2. Corolla tube campanulate, 1.5-2.5 mm long, withered corolla surrounding the 
capsule base before falling a. nee var. gronovii 


a. Cuscuta gronovii var. gronovii. Cuscuta vulgivaga Engelm., Amer. J. Sci. Arts 43:338. 1842. 
Cuscuta gronovii Willd.ex Roem. & Soe var. Vu ke ae marie Trans. Acad: a 
St. Louis 1:508. 1859. Engelmann (1842 it 
specimens for each of the three (as below), but he did not ea which vat ae was typical 
for the species nor did he cite any specimens pertinent to the typical expression of the spe- 
cies, exclusive of varieties. In 1859, he noted for C. gronovii var. vulgivaga that “It is Willdenow’s 
original C. gronovii, in his Hb. nro. 3160” (thus equating var. vulgivaga with var. gronovii). 


202 BRIT.ORG/SIDA 22 


= 


1) 


Cuscuta vulgivaga Engelm. var. [_ | laxiflora Engelm., Amer. J. Sci. Arts 43:338. 1842. LECTOTYPE 
esignated here): U.S.A. ALABAMA: [Oct] 1824, Buckley s.n. (MO). Engelmann cited three (or 

four?) collections: “Western New York on Decodon, Dr. A. Gray; Missouri on Cephalanthus 
and Amphicarpaea, and Georgia, on ? [sic], J. Carey; Alabama, on Salix and Aster, S.B. 
Buckley.” The lectotype sheet has a branch of Salix nigra and one of Symphyotrichum sp 
each hosting Cuscuta 

Cuscuta vulgivaga Engelm. var. [_]glomerata Engelm., Amer. J. Sci. Arts 43:338. 1842. LECTOTYPE 
(designated here): U.S.A. VERMONT: On Leersia, [no date], John Carey s.n. (MO). Engelmann 
noted that “my specimens are from Vermont on Leersia, and New Hampshire, on Solidago, 
both from Mr. J. Carey.” The lectotype specimen has a small branch of the host with Cuscuta 
anda packet with dissected flowers. Mounted on the same sheet is a fragment packet with a 
Carey specimen of Cuscuta “on Solidago” from New Hampshire. It appears likely that both 
specimens are fragments of the ori gi nal collection 

Cuscuta vulgivaga Engelm. var.[_ |tetramera ay hee, Sci. Arts 43:338. 1842. TyPE: U.S.A: 
Engelmann noted only “Connecticut, on Urtica, J. Carey” (HOLOTYPE: MO, presumably, not 
located) 


Pend 


Distribution and ecology.—The most common and widespread dodder in North 
America (Engelmann 1859; Yuncker 1965). CANADA: Alberta, British Colum- 
bia, Manitoba (only a few collections), Ontario, New Brunswick, Nova Scotia, 
Prince Edward Island, Québec, Saskatchewan (only a few collections). U.S.A.: 
Alabama, Arkansas, Arizona, Colorado, Connecticut, District of Columbia, 
Florida, Georgia, lowa, Idaho, Illinois, Indiana, Kansas, Kentucky, Louisiana, 
Maine, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, 
Montana, Nebraska., New Hampshire, New Jersey, New Mexico, New York, North 
Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, Rhode Island, 
South Carolina, South Dakota, Tennessee, Texas, Vermont, Virginia, West Vir- 
ginia, Wisconsin, Wyoming. West Indies (Yuncker 1965) and Europe along wa- 
ter courses (Feinbrun 1972). Growing in a wide variety of climates but appar- 
ently preferring shaded and/or wet environments, sometimes occurring as a 
weed, e.g., in cranberries and praPeviie (Parker & Riches 1993). Flowering Jun- 
Nov. Hosts: ies from hundreds of genera of herbaceous and wood 
plants in various families, including Acanthaceae, Anacardiaceae, Apiaceae, 
Asteraceae, Bignoniaceae, Brassicaceae, Caprifoliaceae, Commelinaceae, 
Convolvulaceae, Cornaceae, Balsaminaceae, Betulaceae, Euphorbiaceae, 
Fabaceae, Lamiaceae, Polygonaceae, Primulaceae, Rosaceae, Rubiaceae, Solan- 
aceae, Urticaceae, Verbenaceae, Vitaceae, and others (see also Gaertner 1950). 
Conservation status: G5 (common) (the same assessment in NatureServe 2005). 


b. Cuscuta gronovii var. latiflora Engelm., Trans. Acad. Sci. St. Louis 1:508. 1859. 
LecTOTYPE (): U.S.A. MISSOURI. Margin of lakes and swamps, in the “American Bottom” oppo- 
site Saint Louis, on Saururus, Sep 1841, Geyer s.n. (MO, designated by Yuncker 1921). Engel- 
mann cited “C. saururi, Engelm!” in synonymy of Cuscuta gronovii var. latiflora. 


] 


Cuscuta saururi Engelm., Amer. J. Sci. Arts 43:339. 1842. Cuscutagronovii Willd. ex Roem & Schult 
var. saururi (Engelm.) MacMill., Metasp. Minnesota Valley 430. 1892. Lecrotyre (Yuncker 
1932): U.S.A. Missourt margin of lakes and swamps, in the “American Bottom” opposite Saint 


COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 203 


Louis, on Saururus, Sep 1841, Geyer s.n. (MO). aa also cited “Alabama, Dr. A. Prout; 
Texas, on Boehmeria, Polygonum, &c., F Lindhe 
Variety latiflora is variable and intermediates with var. gronovii are relatively 
common. In seemingly intermediate specimens, on a single plant flowers can 
be found with the calyx about equaling the corolla tube or somewhat shorter. 
Apparently, the corolla tube may grow to some extent after fertilization occurs, 
thus modifying the ratio between calyx length and corolla tube. In var. gronovii 
all the flowers have the calyx about half the corolla tube length. The size of 
flowers and capsules in var. latiflora is similar to C. cephalanthi, but the latter 
differs in its 4-merous flowers and capsules capped by the withered corollas. 
Forms of var. latiflora with corolla lobes as long as the corolla tube are 
comparable in floral morphology to Cuscuta obtusiflora var. ganas, which 
differs mostly in its depressed-globose capsules with large inter sylar apertures. 
Engelmann (1842) also pointed out that var. latiflora bears some ies with 
C. polygonorum, which is closely similar to C. obtusiflora. 
Distribution and ecology.—The same distribution and hosts as var. gronovit 
but less common. Flowering Sep-Nov. 
Conservation status.—G5 (common) (the same assessment in NatureServe 
2005) 


c. Cuscuta gronovii Var. calyptrata ones Trans. Acad. Sci. St. Louis 1:508. 1859. 
Cuscuta calyptrata (Engelm.) Small, Fl. S.E. U.S. 969. 1903. LectotyPe (Yuncker 1921): U.S.A. 
Western Louisiana, [no datel, J. Gregg s.n. co Engelmann also cited “Texas, Lindheimer!” 

The only differences between var. calyptrata and var. gronovii apparently are 

in corolla size and the way the corolla is shed. In var. calyptrata, as the capsule 

expands, the corolla breaks transversely more or less regularly from the base 
and a permanently narrowed portion of the tube holds the proximal portion of 
the corolla (including the styles and remnants of the stamens) coherent as it is 
lifted atop the capsule. In var. gronovii, the corolla breaks irregularly, trans- 
versely and longitudinally, as the capsule expands and it remains at the cap- 
sule base. In the few plants studied that might be intermediate, the corolla tube 
is not apically narrowed and breaks irregularly but is lifted at the top of the 
capsule at least in some flowers. Variety calyptrata is the most narrowly dis- 
tributed entity of the C. gronovii complex and perhaps the most sharply de- 
fined. However, var. calyptrata is very similar to C. cephalanthi in the unique 

way corolla caps the mature capsule, but the latter differs in commonly 4- 

merous flowers, narrower infrastaminal scales, and dep d-globose capsules. 

The relationships between these three taxa need further study. 

Distribution and ecology.—U.S.A.: Louisiana, Texas. Flowering Aug-Oct. 

Hosts: woody and herbaceous species. 

Conservation status.-T1T2 (critically imperiled to imperiled) (T2, 

NatureServe 2005). 


204 BRIT.ORG/SIDA 22(1) 


2. Cuscuta umbrosa Beyr. ex Hook. Fl. Bor. Amer. 2:78. 1840. (Fig. 1 b, d, g, h). 
Grammica umbrosa (Beyr. ex Hook.) W.A. Weber, Southwest. Naturalist 18:319. 1973. LECTo- 
TYPE (designated here): “Northwestern America,” Douglas s.n. (K). Among three collections, 
Hooker cited “N.W. Coast of America, Douglas.” The sis anaes of C. umbrosa has been 


intertwined with that of C.gronovii var. curta iDelow. ae parent!) obscur Sa ak 
lectotypification of the earlier name from Hool Engelmann (1859) placec 


umbrosa Hook. Lc. in part” in synonymy of C. ere var. curtd, and he cited * Nineuide 
merica, Snes Yuncker (1965) treated Cuscuta gronovii var. curtaasa synonym 

of C. umbrosa but noted only that the type locality of C. umbrosa was “Northwestern America.” 
Cuscuta a see ex Roem. & Schult. var. curta Engelm., Trans. Acad. Sci. St. Louis 1:508. 
(Engelm.) Rydb., Bull. Torrey Bot. Club 40:466. 1913. LecTotyPE (Yuncker 

1921): may America (Fremont'’s 3rd Exped. 79), the type, in the Engelmann Herb. (MO). 


For C. curta (with C. umbrosa as a synonym), Yuncker (1932) lace cited “Northwestern 
America, (Douglas, a specimen in the Kew Herbarium)” as the “type. 
Cuscuta sie hate Rydb., Bull. Torrey Bot. Club 28:501. 1901. LECTOTYPE (formally designated 
here, Reveal in herb. 1977) U.S.A. COLORADO: Cucharas Creek, near La Veta, 7000 ft, 15 Sep 
1900, Vreeland 670 (NY). Rydberg also cited Tweedy 2278, a collection from Sheridan Co. 
Wyoming. 
Stems medium to coarse, yellow to orange. Inflorescence of pedicellate (1-7 mm) 
flowers commonly in dense, paniculate cymose clusters; each flower with one 
ovate to broadly triangular bract at the pedicel base. Flowers 5-merous (some- 
times 4-merous), 2-3.5(-4.4) mm long , not papillose, white-cream, with a few 
to many isolated, round to ovoid translucent laticifers in the calyx, corolla and 
ovary. Calyx campanulate reaching toca. the middle of the corolla tube, divided 
ca. 1/2-2/3 the calyx length, lobes ovate, apex rounded or obtuse, basally over- 
lapping, margins entire to serrulate; corolla tube cam panulate, 1.7-2.3(-2.7) mm, 
lobes mostly ovate to broadly-triangular ovate, rounded-obtuse, 1/3-1/4 the 
tube corolla tube, spreading or reflexed; epicuticular wax absent or reduced to 
a few reticulated rodlets; stamens exserted, anthers on filaments equaling or 
longer than anthers; pollen 3(-4)-zonocolpate, (18-)20-25(-28) um long, 
rounded or truncated at poles, polymorph in the same anther or flower, from 
spheroidal to subprolate (the former shape more frequent); usually tectum 
perforatum to almost microreticulate, puncta 0.4-0.7 um in diameter, rarely 
tectum imperforatum; sexine scabrate with isolated granules; infrastaminal 
scales infrastaminal scales broadly-oblong, and apically truncate to slightly 
bilobed, fringed, 1/2(-1/3)as long as the tube; styles distinct, slender, thickened 
at the base 0.3-0.7(-0.9) mm long; stigmas capitate, globose. Capsules 
subglobose, ovoid to globose-conic or subobpyriform, thickened at the top, 
interstylar oppening relatively small, sometimes with a short and broad but 
discernible neck, 3.5-6.5(-7) X 3-5(-6) mm, indehiscent or irregularly dehis- 
cent. Seeds 3-4 per capsule, (1.8-)2-2.5(-2.8) x 1.5-1.65 mm, dorsoventrally com- 
pressed to obscurely angled, broadly-elliptic to obovate; hilum subrotund, 0.40- 
0.60 x 0.39-0.46 mm, vascular scar of funiculum linear, 0.20-0.25 mm long, 


COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 205 


vertical to slightly oblique. Surface of seed epidermis alveolate when dry and 
papillate, with cells 20-50 tm in diameter. 

Cuscuta umbrosa is similar in some aspects of flower, fruit, and seed mor- 
phology to C. rostrata. Both have infrastaminal scales shorter than the corolla 
tube and large capsules and seeds. Cuscuta rostrata is recognizable by the nar- 
rower infrastaminal scales and especially by the prominently beaked ovary and 
capsule. Cuscuta umbrosa, however, sometimes produces capsules with a short 
and broad neck, which may indicate homology with the beaked capsules of C. 
rostratd. 2n =? 

Distribution and ecology.—CANADA: Alberta., Manitoba, NW Ontario (only 
one collection), Saskatchewan. U.S.A.: Colorado, Kansas, Minnesota, Montana, 
Nebraska, New Mexico, New York (as reported by Mitchell and Tucker 1997; 
possibly a waif), North Dakota, Oklahoma, South Dakota, Utah, Wyoming. Flow- 
ering Jul-Oct. Growing in shaded and/or wet environments. Hosts: species of 
Salix, Ampelopsis, Symphoricarpos, Clematis, Epilobium, Convolvulus, 
Scutellaria, Linum, and other genera. 

Conservation status.—G4G5 (apparently secure to secure) (G5, NatureServe 
2005) 


APPENDIX 1 
VOUCHERS FOR THE sEMsrubpyY 


1a. Cuscuta gronovii var. gronovii (16 collections examined). CANADA. ONTARIO. Leeds Co.: 
Rear of Leeds Twp., 1 km N of Oak Leaf, 6 km SE of Delta, 44 35'20"N 76 02'55"N, 3 Sep 1990, 
Catling 5111 (DAO). NOVA SCOTIA. Lunenburg Co.: Wentzell Lake, 17 Aug 1921, Fernald & Long 
a (NY). U.S.A. ALABAMA. Montgomery Co.: Alabama River Bottoms, 4 mi N of Montgomery, 
1 Oct 1943, Kpeooer et al.s.n. (NY). ARKANSAS. Hot Springs Co.: Magnet Cove, 470 ft, 2 Oct 1938, 
ae 2062 (NY). FLORIDA. Taylor Co.: vicinity of Adams Beach, 16 Oct 1961 Peace 61565 
(NY).GEORGIA. Fulton Co.: Atlanta, 1164 Clifton Rd., 1 Oct 1956, Burbanck s.n. (NY). INDIANA. Fran- 
klin Co.: bank of river 4 mi S of Laurel, 12 Oct 1924, Deam 41026 (NY). MAINE. Knox a seashore 
polveuas 14 Aug 1952,Friesners.n. (NY). MASSACHUSETTS. Barnstable Co.: 26 Aug 1928, Fernald 
ieee Grayane 480 (NY). MICHIGAN. Kent Co.: 7 mi N of Lowell, 28 Aug 1931, Yuncker & 
oe 3792 (NY). MISSISSIPPI. Noxubee Co.: 3 mi of Macon, 22 Oct 1960, McDaniel 2276 (NY). MIS- 
SOURI. Jefferson Co.: 6.5 mi W of DeSoto, 38 07’30"N 90 40'30" , 25 Aug 1991, Raven & Raven 
27930 ee Saint Louis Co.: near Saint Louis, 1878, Eggert s.n. (NY). NEW JERSEY. Middlesex Co.: 
opp w Brunswick, Oct 1918, Mackenzie s.n. (NY). OHIO. Ottawa Co.: Turtle Creek, Ottawa 
eer tet Refuge, 570 ft, 22 Aug 1996, Hill 28542 (NY). PENNSYLVANIA. York Co.: near McCall's 
Ferry, 9 Sep 1893, Heller & Halbach 1357 (NY). WISCONSIN. Waushara Co.: 2.5 mi SSW of Borth, 15 
g 1984, Bewick 100 
1b. Cuscuta gronovii var. latiflora (12 collections examined). U.S.A. INDIANA. Daviess Co.: 
1.5 mi SE of Newberry, 21 Sep 1932, Deam 53259 (NY). Pasey Co.: 12 mi SW of Mt. Vernon, 20 Aug 
1922, Deam 37712 (NY). Sullivan Co.: along Busseron Creek, 2 mi E of Sullivan, 3 Oct 1931, Deam 
51395 (NY). IOWA. Wapello Co.: Cliffland, NW1/4 Sec. 11,T-71N, R-13W, 30 Sep 1954, Davidson 4051 
(NY). MARYLAND. (no county given): at mouth of Little Gunpowder River, 2 Sep 1902, Shull 289 
(NY). MICHIGAN. Kalamazoo Co.: Portage Twp., 20 Jul 1936, Hanes s.n. (NY). NEW JERSEY. Mercer 
Co.: Duck Island, below Trenton, 21 Aug 1921, Mackenzie s.n. (NY). NEW YORK. Nassau Co.: Long 


= 
= 


206 BRIT.ORG/SIDA 22(1) 


Island, Oyster Bay, 11 Sep 1926, Ferguson 6091 (NY). Suffolk Co.: Long Island, Water Mill, 10 Aug 1925, 
Ferguson 4293 (NY). Wayne Co.: Lake Ontario, Fairbanks Point, Williamson Twp., 246 ft, 10 Aug 1939, 
Hinkey 4484 (NY). TEXAS. Wood Co.: Lake Ellis, 3 Sep 1942, Lundell 11750 (NY). VIRGINIA. Princess 
Anne Co.: along North Landing River, near Creed’s, 9 Sep 1935, Fernald et al. 5011 (NY). 

1c. Cuscuta gronovii var. calyptrata (1 collection examined). U.S.A. TEXAS. Rockwall Co.: 
3.25 mi NW of Rockwall, 22 Oct 1946, Cory 52529 (SMU) 


2. Cuscuta umbrosa (14 collections examined). CANADA. ALBERTA: Lethbridge, 30 Aug 
1971, Allen 150 (DAO). MANITOBA: Aweme (?), 9 Aug (no year), Criddle s.n. (DAO); Delta, 
S13-T14-R7, 20 Aug 1953, Love & Léve 6/99 (DAO); Delta Marsh, veers Field Station, 
S50°L17N 98° 24°W, 9 Aug 1984, Szumigalski 18 (DAO); Otterburne, 13 A 954, Bernard 497 
(DAO); Winnipeg, Mulvey Avenue, Sep 1975, Fields s.n. (DAO). ona Carnduff, 
20 Aug 1948, Budd & Lodge 1197 (DAO); 10 mi S of- and 0.5—1 mi E of Gainsborough, Hutler 
River Valley, NE1/4 Sec. 9 Tl R30 W1, 29 Jul 1988, Harms 39594 (DAO). U.S.A. COLO- 
RADO. Teller Co. (?).: Manitou oe eae Forest?), 31 Aug 1892, Mulford s.n. (NY). MIN- 
NESOTA. Carlton Co.: 3 mi SW ond du Lac, 22 Aug 1945, Moore & Huff 18199 (DAO). 
NORTH DAKOTA. Cass Co.: 2 9 Sep 1950, Stevens 1228 (DAO). SOUTH DAKOTA. 
Potter Co.: Forest ene ae 1892, Griffiths & Glosser s.n. (NY). UTAH. Salt Lake Co.: City 
Creek Canyon, 8000 ft, 10 Aug 1883, Leonard 250 (NY). 


ls 


ACKNOWLEDGMENTS 


— 


Dan Austin, Lytton Musselman and an anonymous reviewer provided valuable 
comments and suggestions that improved an earlier version of the manuscript. 
We thank directors and/or curators from ACAD, ALTA, ARIZ, ASU, BRIT, DAO, 
FE GH, HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, 
RSA, SASK, SFS, TEX @ LL, TUP UBC, UC &JEPS, UNB, UNM, US, USAS, UWO, 
UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Special apprecia- 
tion is extended to NY staff for approving and preparing the numerous volumi- 
nous loans. Robert Vogt and Meine Jutta (B) provided helpful information re- 
garding the type collection of Cuscuta gronovii, and Kaj Vollesen (K) provided 
information regarding the type collection of Cuscuta umbrosa. Alexandra Smith 
assisted us with th lectron i d Elma Schweigert checked 
the translation of the Spanish abstract. 


REFERENCES 


Coste, M., G.L. Nesom, and S. Steranovic. 2006. Taxonomy of the Cuscuta pentagona com- 
plex (subsect. Arvenses: Convolvulaceae) in North America. Sida 22:151-175. 

Cains, WJ.and B.A. Foro. 1988. The parasitic dodders (Cuscuta: Cuscutaceae) in Ontario. 
Canad. Field-Nat. 102: 209-215. 

DareysHire, S.J. 2003. Inventory of Canadian agricultural weeds. Agriculture and Agri-Food 
Canada, Research Branch, Ottawa, ON. 396 pp.Electronic Publication: <http://res2.agr.ca/ 
ecorc/weeds_herbes/title-titre_ehtm>. 

Dawson, J.H. 1984.A vegetative character that separates species of Cuscuta.Proc. 3rd Interntl. 
Symposium on Parasitic Weeds. Pp.184-187 


COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 207 


ENGELMANN, G. 1842. A monograph of the North American Cuscutineae. Amer. J. Sci. 43: 
333-346 

ENGELMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with 
critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 
1:453-523. 

Feinsrun, N. 1972. Cuscuta L.In:T.G.Tutin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H.Valen- 
tine, S.M.Walters, D.A.Webb, eds. Flora Europaea, Vol. 3, Diapensiaceae to Myoporaceae. 
Cambridge University Press. Pp. 74-77. 

FOGELBERG, S.O. 1938. The cytology of Cuscuta. Bull. Torrey Bot. Club. 65:631-645. 

Gaertner, E.E. 1950. Studies of seed germination, seed identification, and host relation- 
ships in Dodders, Cuscuta spp. Mem. Cornell Agric. Exp. Sta. 294:1-56. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the 
vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T. 
and C.A. Meacham. 1999. Synthesis of the North American Flora, Version 1.0. North 
Carolina Botanical Garden, Chapel Hill, NC. 

MitcHett, R.S.and G.C. Tucker. 1997.A checklist of New York state plants (rev. ed.). New York 
State Bull. 490. New York State Museum, Albany. 

NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web application, 
Version 4.0. NatureServe, Arlington, Virginia. <http:/ ve.org/expl 
(Accessed: November 16, 2005) 

Parker, C.and C.R.RicHes. 1993.Parasitic weeds of the world. Biology and control. CAB Inter- 
national, Wallingford, UK. 

USDA, NRCS. 2004. The PLANTS database. National Plant Data Center, Baton Rouge, LA. 
Electronic Publication, Version 3.5: <http://plants.usda.gov 

YUNCKER, T.G. 1921. Revision of the North American and West Indian species of Cuscuta. 
Illinois Biol. Monogr. 6:91—231.Reprinted 1970, Johnson Reprint Company, N.Y. 

Yuncker, T.G. 1932.The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. 

Yuncker, 1.G. 1943. Nomenclatural changes in the genus Cuscuta and notes on some . 
American species. Bull. Torrey Bot. Club 70:61-67. 

YuNcKER, 1.G. 1965. Cuscuta. North American Flora, ser. 2,4 


at 


oe 


208 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Vinaya GuaTE, Hema Sang, and S.S. RANADE (compilers and editors). 2004. Focus 
on Sacred Groves & Ethnobotany. (ISBN 81-8540502-6, hbk.). Proceedings 
of the National Seminar on Ethnobotany & Sacred Groves: “Role in Con- 
servation Strategy for India.” [Edited by Dr. Vinaya Ghate] Prism Publica- 
tions, Chembur, Mumbai (India). Distributor: Timely Management 
Consl Ser.Put.Ltd., 14/4 Shivpuri, Sion Trombay Road, Chembur, Mumbai 
400 071, INDIA. (Orders: PH: 91-22-2522 7616 / 2529 5725 Fax: 91-22-2524 
2484, ajitn@vsnl.com). Rs.900 (US $50), 253 pp., photographs, 7.5" x 10" (19 
x 25cm). 


Sacred groves are re ae! small, some no more than a stand of trees, few larger than an acre of 
mixed climax vegetation. Within them the locals worship the deities to whom the groves are dedi- 
cated and also hold festivities of dance and song in their honor. Because collecting live or fallen plant 
materials or the cutting of them is taboo, the groves may be the last hold out of plants that could be 
threatened with extinction as the surrounding forests are cleared for human purposes. The groves 
are therefore important for conservation. 

D. Vaktar, who taught at the University of Pune, was a leader in the study of ethnobotany 
of western India. The conference and this book commemorate his work. The first part of the book 
contains articles that Dr. Vaktar (and co-authors) published between 1973 and 1998. Part 2 comprises 


papers presented at the seminar: studies of botany of various locales, plants important in ie cau 
observances, medicinal plants and their applications. The need for conservation of plants in the sa- 
cred groves and elsewhere is obvious in all the p ee Karges (retired TCUL a Botani- 
cal an Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(1): 208. 2006 


TAXONOMY OF THE CUSCUTA INDECORA 
(CONVOLVULACEAE) COMPLEX IN NORTH AMERICA 


Mihai Costea (corresponding author) Guy L.Nesom 
Department of Biology Botanical Research Institute of Texas 
Wilfrid Laurier University 09 Pecan Street 
75 ae Avenue West Fort Worth, Texas 76102-4060, U.S.A. 
Wate ntario gnesom@brit.org 


N2L ca CANADA 
mcostea@wlu.ca 


Sasa Stefanovic 


Department of Biology 
University of Toronto at Mississauga 
3359 Mississauga Road 
Mississauga, Ontario 

L5L 1C6, CANADA 


ABSTRACT 


The Cuscuta indecora complex (Cuscuta subsect. Indecorae) in North America is characterized by 
fleshy, papillose flowers, corolla lobes with inflexed ae and capsules with a pmeccned styl one 
dium. It includes C. indecora, C. coryli, C. warneri, and C. jepsonii. Recent 
to treat the latter three taxa at varietal rank within C. indecora or of it. We maintain C. 
warneri and C. jepsonii at specific rank because of their distinctive morphology, even Aves the 
latter is known only from the type collection and the former froma very limited number of speci- 
mens. Cuscuta indecora includes var. indecora, var. longisepala, and C. indecora var. attenuata (Wa- 
terfall) Costea, comb. et stat. nov. Cuscuta indecora vars. bifida and neuropetala are treated as syn- 
.indecora var. indecora. Cuscuta warneriand C. coryliare reported as new floristic records 


from New Mexico and Québec respectively. 


RESUMEN 


El complejo Cuscuta indecora (Cuscuta subsect. Indecorae) en América del Norte esta caracteriza 

por flores papilosas y suculentas, lobulos de la corola con ae inflexionadas, ot ieee: con un 
estilopodio engrosado. El complejo incluye C. indecora, C. coryli, C. warneri, y C. jepsonii. 
Recientemente algunos taxonomos ee Dae tratar las me tres ee dentro de la 
categoria C. indec jue C. warneriy C. jepsonii estan 
en una categoria especial por su oo a a pesar de que la Ace se conoce sdlo por la 
coleccion del tipo, y la primera es conocida mediante un numero de especuenes eye unease: 
Cuscuta indecora incluye var. indecora, var. nies y C. indec ostea 

b. et stat. nov. Cuscuta indecora vars. baa neuropetala son tratadas cé indénim 

indecora var. indecora. Cuscuta warneri y C. coryli figuran como nuevas citas floristicas de Nueve 


uUala 


México y Quebec, respectivamente 


Cuscuta subsect. Indecorae Yuncker is characterized by fleshy, papillose flow- 
ers, corolla lobes with inflexed tips, and capsules with a thickened stylopodium 


SIDA 22(1): 209 — 225. 2006 


210 BRIT.ORG/SIDA 22(1 


(Yuncker 1932, 1965). In Yuncker’s view, it included three North American spe- 
cies (C. indecora Choisy, C. coryli Engelm., and C. warneri Yuncker) and one 
from South America (C. stenolepis Engelm.). A fifth species, C. attenuata Wa- 
terfall, was added to subsect. Indecorae by Prather and Tyrl (1995), although 
Waterfall (1971) had suggested that C. attenuata resembled C. compacta Juss. of 
subsect. Lepidanche Engelm. Cuscuta attenuata is closely similar to C. indecora, 
but Prather and Tyrl (1995) maintained both at specific rank, emphasizing their 
apparent reproductive isolation. Cuscuta warneri was treated by Beliz in her 
Ph.D. thesis (1986) as a variety of C. indecora; this unpublished combination 
has been included in some recent North American overviews (e.g., Kartesz 1999). 
Cuscuta jepsonii Yuncker, which was initially treated in subsect. Californicae 
Yuncker (Yuncker 1932), has been recently considered conspecific with C. 
indecora (Beliz 1993, 2002). The taxonomic status and relationships of these 
taxa are reevaluated here, based on morphology and micromorphology of flow- 
ers, capsules, seeds, and pollen. 


METHODS 


Descriptions of morphology are based on samples from specimens from NY, 
which includes Yuncker’s herbarium (Appendix 1). Measurements and pictures 
were taken witha scanning electron microscope Hitachi $-570 at 15 KV. Samples 
were coated with 30 nm gold using an Emitech K 550 sputter coater. Terminol- 
ogy regarding the micromorphology of flowers, seeds and capsules, and pollen 
were described in detail in the first paper published in this issue (Costea et al. 
2006). Conservation status was assessed using NatureServe (2005) ranks and 
criteria. 


TAXONOMY 


Cuscuta indecora and C. coryli.—Cuscuta indecora is a highly variable and 
common species both in North America and South America (Engelmann 1859; 
Yuncker 1921, 1932, 1965; Beliz 1986; Prather & Tyrl 1995). Cuscuta coryli is less 
variable and is sympatric with C. indecora over a significant geographic area 
(see below). Yuncker (1932) mentioned that C. coryli “is closely related to C. 
indecora, but is distinguished by its often 4-parted flowers which are commonly 
smaller (ca. 2 mm long), rudimentary scales and shape of fruit.” We find that 
flowers of C.coryli may be both 4-merous and 5-merous on the same specimen, 
and sometimes even 3-merous. Engelmann (1843) noted “flowers frequently 5- 
parted” for C. coryli, but he later 1859) modified this observation to flowers 
“mostly 4-parted.” Although the type and many collections of C. coryli appar- 
ently have more 4-merous than 5-merous flowers, 5-merous flowers predomi- 
nate in some specimens. Ranges of variation in floral size in C. indecora and C. 
coryli are closely similar. Cuscuta indecora has small flowers, 2.1-2.7 mm long, 


COSTEA ET Al OF CUSCUTA INDECORA COMPLEX 211 


with the corolla tube 0.9-1.5 mm long; flowers of C. coryli are 1.7-2.6(-3) mm 
long, with the corolla tube (0.7-)1-L.4 mm long. 

Such observations perhaps led Beliz (1987-1988, in herb. NY, GH) to con- 
sider Cuscuta coryli conspecific with C. indecora. Although they are clearly re- 
lated and sometimes overlapping in several character states, the two taxa can 
be usually distinguished using a combination of characters (see key and de- 
scriptions below) and their recognition as separate species is appropriate. 

Description of infrastaminal scales in these two species has been confus- 
ing. For example, in Cuscuta coryli, Engelmann was consistent, describing the 
infrastaminal scales as “appressed, bifid, consisting of a few teeth ... one or two 
teeth on each side of the filament (1942) ... or lobes laterally adhering to the 
lower (attached) part of the filament” (1859). Yuncker (1921, 1932) added “scales 
rudimentary, bifid, toothed, ordinarily reduced to toothed wings on either side 
of filament attachment.” Figure 42e (1921) depicted one bifid infrastaminal scale 
with | or 2 fimbria on the each side of the staminal filament, while in Fig. 39 
(1932), these lateral fimbria are replaced by dentate lobes (wings). In his 1965 
treatment, Yuncker removed the term “bifid” from the description of 
infrastaminal scales of C.coryliand applied it to C. indecora var. bifida Yuncker, 
which was described as identical in other respects to C. indecora var. indecora. 
We find that the infrastaminal scales of var. bifida are not truly bifid, but rather 
the spathulate scales may have 2 or 3(-4) deeper apical incisions, which create 
2 or 3(-4) lobes that are further fringed. “Normal” scales may occur in the same 
{lower together with lobed ones. Such plants are regarded here as populational 
variants of C. indecora var. indecora. Bifid scales, as pointed out by Engelmann 
(1842, 1859), are characteristic of C. coryli. 

The varieties of C. indecora and the status of Cuscuta attenuata.— Yuncker 
(1965) treated Cuscuta indecora with three varieties (var. indecora, var. bifida, 
and var. longisepala): as noted below, we tentatively maintain var. longisepala 
but var. bifida is not appropriately recognized. Cuscuta indecora var. neuropetala 
(Engelm.) Hitchc., distinguished by its relatively larger flowers, has been ac- 
cepted in recent overviews of the genus (Beliz 1993, 2002; Kartesz 1999). We 
find that although var. neuropetala can be often identified, its connection to var. 
indecora by a series of intermediates makes taxonomic recognition unfeasible. 

Prather and Tyrl (1995) observed that Cuscuta attenuata is morphologically 
similar to C. indecora. Indeed, Yuncker annotated (in herb. NY) some collec- 
tions of C. attenuataas C. indecora var. longisepala. The present study substan- 
tiates the similarities, which extend to the morphology and micromorphology 
et Epona seeds, pollen, and capsules. The geographic range of C. attenuata 

p within that of var. indecora, but preliminary evidence suggests 
that C. attenuata may have a narrower host preference. In UPGMA and princi- 
pal components analyses by Prather and Tyrl, C. attenuata is part of a single 


212 BRIT.ORG/SIDA 22(1) 


cluster with C. indecora var. indecora and var. longisepala; univariate analyses 
separated these three taxa, although they were broadly overlapping in morphol- 
ogy. Prather and Tyrl (p. 456) concluded that “C. attenuata is a distinct species 
albeit morphologically similar to C. indecora. In the absence of reproductive 
isolation we might treat C. attenuata asa variety of C. indecora.” Experimental 
crosses by Prather and Tyrl between C. attenuata and both varieties of C. 
indecora produced neither fruits nor seeds, while populations of C. attenuata 
were interfertile. Whether var. indecora and var. longisepala were interfertile 
was not reported. 

For consistency with the degree of morphological difference between other 
Cuscuta species (as we are recognizing them), C. attenuata is treated here at 
varietal rank within C. indecora. In view of its apparent reproductive isolation 
and host specialization, it isa more strongly defined entity than var. longisepala, 
which also is broadly sympatric with var. indecora. Var. longise pala is tentatively 
maintained here, until its biology and evolution may be better understood. 

Cuscuta warneri.—T his species is a strikingly distinct dodder. Each calyx 
lobe is apically prolonged into a conical spur-like projection (Fig. 4 a,b,d), 
infrastaminal scales are oblong with truncate and dentate apex, and capsules 
have a collar-like apex and very short styles (Fig. +d). The calyx spurs are 
accrescent, relatively small in flower and reaching maximum size in fruit, when 
they detach easily (at least on dry material). Cuscuta warneri shows strong simi- 
larity to C. indecora in morphology and micromorphology of perianth, cap- 
sules, seeds, and pollen (as also observed by Yuncker 1960). Beliz 1986) wrote 
that “critical studies, however, indicate that C. warneri is probably an abnormal 
specimen of C. indecora ... and until more material is available for studies, C. 
warneri is recognized asa variety of C.indecora.” Our observations, in contrast, 
do not indicate that C. warnertis teratological. Flowers have all the components 
and they are fertile, each capsule usually with 2 seeds in which the embryos 
appear to develop normally. The fact that C. warneri is known only from the 
type locality and a collection from New Mexico (see bellow) indicates that its 
distribution is localized, but its morphological distinction justifies continuing 
recognition at specific rank. 

The spur-like projections of the perianth of Cuscuta warneri have an un- 
known biological role. Similar morphology also is encountered in other, more 
distantly related species: C. runyonii Yuncker, C. applanata Engelm., C. 
boldinghii Urban, and C. chapalana Yuncker. 

Cuscuta jepsonii.—T his species was described and included by Yuncker 
(1921, 1933) in subsect. Californicae because infrastaminal scales are absent or 
reduced to ridges. Beliz (1986) initially considered it to be a synonym of C. 
californica Hook. & Arn. var. papillosa Yuncker, but she later (1993) treated it 
as asynonym of C. indecora var. indecora. She did not provide substantiating 
evidence for these decisions. The species is known only from the UC holotype. 


sy 


COSTEA ET AL., TAXONOMY OF CUSCUTA INDECORA COMPLEX 213 


Based on study of this collection, we confirm that C. jepsonii may belong to 
subsect. Indecorae, where it represents an extreme case of infrastaminal scale 
reduction. The 5-merous flowers and papillae morphology are similar to C. 
indecora, from which it differs by very small anthers, 0.2-0.3 mm long, which 
are more like those of C. coryli. 

Infrastaminal scales vary to some extent in many Cuscuta species. Never- 
theless, we know of no instance of complete reduction of infrastaminal scales 
in a species where scale development is characteristically normal, even if vari- 
able. For this reason, and until additional material can be studied, C. jepsonii is 
maintained as a distinct species. 


KEY TO THE SPECIES OF CUSCUTA SUBSECT. INDECORAE IN NORTH AMERICA 
1. Infrastaminal scales absent or reduced to ridges 4.C. jepsonii 
1. Infrastaminal scales present 
2. Calyx lobes with an apical spur-like projection; infrastaminal scales dentate at 
3 


.C. warneri 


apex 
2. Calyx lobes without spur-like projections; infrastaminal scales fimbriate. 

3, Flowers commonly 5-merous, 2-5.3 mm long, infrastaminal scales united with 
the corolla tube for 1/3-1/2 (rarely 3/4) of their length, subspathulate to 
spathulate, rarely 2-3 lobed, apex rounded rarely truncate, with (6-)20-35(- 
50) fimbria; capsule yellowish and + Ue nena. when dried, globose, 
subglobose to slightly depressed-globose, 0.8-1.5 times wider than long, the 
suture lines between the 2 ene not or ony slighty cleiecce 1.C.indecora 

3. Flowers (3-)4—5-merous, 1.7-2.6(-3 ited wit 
the corolla tube for most of their length, oblong, bifid with 1-3 ee on 
each side of filament attachment, rarely + truncate with 3-6 fimbria; capsule 
brown and not semi-transparent when dried, initially globose, becomes 
depressed, 1.6-2.4 times wider than long, the suture lines between the 2 
carpels depressed, forming a longitudinal groove on the opposed sides of 
capsule 2.C. coryli 


1. cmc indecora cee Men Soc. Phys. Genéve 9:278, t.3, £.5. 1842 (Fig. 1, 
a,d 1&8, h).c 1 Choisy ex Engelm. var. indecora (Choisy) Engelm , Trans. 
ie fe Sci. St. Louis 1:502. 1859. Grammica indecora (Choisy) W.A. Weber, Southw. Naturalist 
18:319. 1973. Epithymum indecorum (Choisy) Nieuwl. & Lunell, Amer. Mid]. Naturalist 4:511. 
1916. TyPE: MEXICO. [TAMAULIPAS]: “Mexicum ad Matamoros,” Berlandier 2285-865 (HOLO- 
TYPE: G-DC; ISOTY PES: MO, P) 


Stems 0.4-0.7 mm thick, yellow to orange. Inflorescences loose to dense, pan- 
iculate-cymose clusters, sometimes originating endogenously; pedicels 0.5-6 
mm long, papillate-hispid to glabrous; bracts one (rarely 0) at the pedicel base, 
ovate to lanceolate. Flowers (4-)5-merous, 2-5.3 mm long fleshy, translucent- 
white when fresh, of the same color or dark-brownish when dried; epidermal 
cells arranged in rows, fleshy, with anticlinal walls convex (dome-like), addi- 
tionally with cylindric-conical papillae 40-80 um long; when flowers dry up, 
rows of dehydrated epidermal cells are usually easily discernible; epicuticular 
wax represented by longitudinally reticulated rodlets; laticifers isolated or in 


214 BRIT.ORG/SIDA 22(1) 


Fic.1. Floral 


. | j lah 
bar 75 Lum) g var ong sepa a; var 


C. indecora: a—e. var. indecora (scale bar = 1 mm); f. Papillae on the calyx of var. indecora (scale 


ta ferala | 


COSTEA ET AL., TAXONOMY OF CUSCUTA INDECORA COMPLEX 215 


longitudinal rows, ovoidal to elongated, present in the perianth along the 
midveins, ovary and capsules often running longitudinally and yellowish-or- 
ange colored. Calyx cupulate, 1/2 tosomewhat longer than the corolla tube, di- 
vided 1/2-2/3 of the length, lobes triangular-ovate to lanceolate, acute to at- 
tenuate, more or less overlapping at the base. Corolla tube campanulate, 
campanulate-cylindric, subglobose or suburceolate, 1-3.2 mm long, lobes 0.7- 
1.5mm long triangular-ovate, acute, ca. 1/3 to equaling corolla length, suberect 
to erect, apically inflexed. Stamens barely exserted or enclosed; anthers elliptic 
to oblong, (0.5-)0.6-0.9(-1.2) x 0.3-0.5 mm long; filaments equaling or longer 
than anthers; pollen 3(-+)-zonocolpate, subprolate to prolate, 24-36 um long, 
tectum imperforatum or with a few isolate puncta, sexine scabrate with iso- 
lated granules. Infrastaminal scales reaching the filaments, united with the 
corolla tube for 1/3-1/2 (rarely 3/4) of their length; subspathulate to spatulate, 
apex rounded, rarely truncate or 2-3(-4) lobed, with (6-)20-35(-50) fimbria. 
Styles distinct, 1-2.5 mm long, + unequal, evenly filiform, suberect or weakly 
divergent; stigmas capitate, globose. Capsules yellowish to light-brown and + 
semi-transparent when dried (pericarp thin), glabrous, subglobose, globose, to 
slightly depressed-globose, 0.9-1.5 wider than long, narrowed and thickened 
around the style bases, indehiscent or irregularly dehiscent, surrounded or 
capped by the withered corolla; pericarp epidermis smooth. Seeds 2-4 per cap- 
sule, 1.42-1.86 x 1.25-1.6 mm, shape heterogeneous on the same plant: dorsoven- 
trally compressed to weakly angled, broadly elliptic to transversely oblique, 
hilum subterminal, rarely almost terminal, broadly elliptic, 0.40-0.45 x 0.32- 
0.36 mm: vascular scar linear, 0.15-0.18 mm, vertical; seed surface variable: a) 
epidermis cells more or less polygonal and puzzle-like, b) alveolate when dry 
and papillose when wet, and c) only some cells are papillose and the rest are + 
puzzle-like; size of epidermal cells 20-50 um in diameter. 


KEY TO THE VARIETIES OF CUSCUTA INDECORA 


1. Calyx lobes ovate triangular, reaching ca. 1/2 of the corolla tube a.C.indecora 
var.indecora 
1. Calyx lobes lanceolate mostly longer than the corolla tube. 
2. Calyx lobes acute; flowers in loose clusters; parasitic on a wide range of species 
b. C. indecora var. longisepala 
2. Calyx lobes attenuate; flowers in dense clusters; parasitic primarily on /va annua 
c.C. indecora var. attenuata 


a. Cuscuta indecora var. indecora. (Fig. 1 a,b,c,d,e). Cuscuta neuropetala Engelm., Amer. 
J. Sei. Arts 45:75. 1843. Cuscuta indecora Choisy var. neuropetala (Engelm.) Hitche., Contr. 
US. Natl. Herb. 3:549. 1896. TyPE: U.S.A. TEXAS. [Harris Co.|: in wet prairies near Houston, on 
pons suchas Liatris, Solidago, Helianthus, oe and on ‘ifera 

43, Lindheimer 124 (HOLOTYPE: MO; ISOTYPES: NY, 


Cuscuta neuropetala Engelm. var. littoralis Engelm., Boston J. Nat. Hist. 5:223. 1845. LECTOTYPE 
(here designated; Beliz 1986, in herb.): U.S.A. TEXAS. [Galveston Co.]|: Galveston, Apr 184 


216 BRIT.ORG/SIDA 22(1) 


Lindheimers.n. ae ISOLECTOTYPE: NY). The protologue noted “Seashore of onsen Island, 
on Lycium ca 1 Borrichia frutescens, Iva frutescens, etc. Flowers in 

Cuscuta ee a var. hispidula Engelm., Amer. J. Sci. Arts 43:341. 1842. ee taindecora 
var. his (E ) Yuncker, Illinois Biol. Monogr. 6:148. 1921. Type: U.S.A. TEXAS. [Harris 


Co.| “in ao ne ster at prairies, west of Houston, on Euthamia, Schrankia, Aster, Ambrosia, 


Evolvulus, and other low herbs, flowering in April and May, FE Lindheimer’ s.n. (HOLOTYPE: 
MO, presumably). Yuncker (1921) cited Berlandier 2285-MO as the type of C. indecora var. 
hispidula, but that collection is interpreted here as the holotype of C. indecora (var. indecora). 
Cuscuta hispidula Engelm., Amer. J. Sci. Arts 45:75. 1843. Type: U.S.A. TEXAS. [Harris Co.]: Engel- 
mann did not cite a collector but noted “in dry and sterile prairies west of Houston. Flower- 
ing in April and May,” apparently referring to the same Lindheimer collection typifying C. 
verrucosa var. hispidula (above). Cuscuta hispidula apparently was not intended by Engel- 


mann to be a new combination based on the earlier C. verrucosa Engelm. var. hispidula, as he 


wrote “Cuscuta hispidula n. sp.” and “Compare the remarks made in Vol. XLIII p. 341, under C. 
verrucosd.” 

Cuscuta indecora var. bifida Yuncker, Illinois Biol. Monogr. 6:149. 1921. LECTOTYPE ah es 
here): U.S.A. NEVADA. Twin Springs, May-Oct, Purpus 6343 (UC 124538). Two sheets at UC a 
marked as “isotype” of var. bifida: the other is Purpus s.n., without date (UC 124541). 

ogy.—CANADA: Saskatchewan. U.S.A.: Alabama, Arkan- 

sas, Arizona, California, Colorado, Connecticut, Florida, Georgia, Idaho, lowa, 

Illinois, Kansas, Kentucky, Louisiana, Maryland, Michigan, Minnesota, Missis- 

sippi, Missouri, Montana, Nebraska, Nevada, New Jersey, New Mexico, North 

Carolina, North Dakota, Oklahoma, South Carolina, South Dakota, Tennessee, 

Texas, Utah, Virginia, Washington, West Virginia, Wyoming. MEXICO; WEST 

INDIES; SOUTH AMERICA. It is probably the third most common dodder in 

North America, after C. gronovii and C. campestris. It is listed as a “locally” im- 

portant weed in the U.S.A. and Argentina (Parker & Riches 1993). Flowering 

Jul-Nov. Hosts: wide range of herbaceous and woody species, e.g., Agalinis, 

Baccharis, Borrichia, Chenopodium, Eupatorium, Helianthus, Heterotheca, Hy- 

pericum, Ipomoea, Iva, Kosteletzkya, Lepidium, Ligustrum, Myrica, Pluchea, 

Polygonum, Rhynchosia, Solidago, Suaeda, Symphyotrichum, Tephrosia, Vernonia. 

Conservation status.—G5 (common) (NatureServe 2005). n = 15 (Pinkava et 

al. 1974); 2n = 30 (Pazy & Plitmann 1995). 


— 


Distribution and eco 


Cou 


—/ 


b. Cuscuta indecora var. longisepala Yuncker, [linois Biol. Monogr. 6:149, Fig, 
44, 97.192] (Fig. 1g). Type: US.A. TEXAS. On the Blanco, Wright s.n. (HOLOTYPE: MO), 
Distribution and ecology.—U.S.A.: Texas. MEXICO; SOUTH AMERICA. Flower- 
ing summer-fall. Hosts: herbaceous and woody species. Conservation status: 
T2T1 (imperiled to critically imperiled) (not yet assessed by NatureServe 2005). 
c. Cuscuta indecora var. attenuata (Waterfall) te comb. & stat. nov. (Fig. 


Cuscuta attenuata Waterfall, Rhodora 73:575. 1971. Type: U.S.A. OKLAHOMA. McCurtain 
Co.: Waterfall Creek, 8 mi S and 2 mi E of Idabel, aus 17157 (HOLOTYPE: OKLA; ISOTYPE: 
sH) 


eT 


Distribution and eco 
1993 


ogy.—U.S.A.: Kansas, Oklahoma, Texas (Prather & Tyrl 


COSTEA ET AL., TAXONOMY OF CUSCUTA INDECORA COMPLEX 217 


Conservation status.—imperiled (G2) (Natureserve 2004). Flowering late 
Aug-Oct. Hosts: Iva annua, rarely Symphyotrichum spp., mudflats, floodplains, 
and disturbed areas. 

Conservation status.—T2T1 (imperiled to critically imperiled) (G2, 
Natureserve 2005). 2n = 30 (Prather & Tyrl 1993). 


2. Cuscuta coryli Engelm., Amer. J. Sci. Arts 43:337. 1842. (Fig. 2, Fig. 3 b, c, D) 
TyPE: U.S.A. MISSOURI [St. Louis Co.|: “on Corylus near St. Louis,” Sep 1841, Engelmann s.n. 
(HOLOTYPE: MO; ISOTYPE: GH). The protologue reads “on Corylus, in the barrens west of St. 
Louis, in August and September.” 

Cuscuta coryli Engelm. var. stylosa Engelm., Amer. J. Sci. 43:337. 1842. TYPE: U.S.A. Missourt. [St. 
Col “on Solidago, St. Louis,” Sep 1841, Engelmann s.n. (HOLOTYPE: MO; ISOTYPES: GH, US). 
The protologue reads “On Solidago, in dry prairies near St. Louis.” 


nom. invalid.| Appare ntly a renaming 


Cuscuta inflexa Engelm., Trans. Acad. St. Louis 502.1859 
5 : 


uscutd coryli, as the latter Was cited immediately and first in synonymy (other names at 
specific rank also were cited). A number of collections were cited in the protologue. 


Inf] lly 4 


Stems 0.30-0.50 mm thick, yellow to orange 
times loose), paniculate-cym lusters, sometimes arenere eidererouely 
pedicels 0.5-3 mm long, glabrous; bracts one at the pedicel base, ovate to lan- 
ceolate. Flowers 4-5-merous (rarely 3-merous), 1.7-2.6(-3) mm long, fleshy, 
white when fresh, commonly dark-brownish when dried; epidermal cells fleshy, 
organized in rows with anticlinal walls convex (dome-like); papillae like those 
described in C. indecora usually absent; when flowers dry up, rows of dehy- 
drated epidermal cells are more or less inconspicuous; epicuticular wax present 
represented by longitudinally reticulated rodlets; laticifers isolated or in longi- 
tudinal rows, ovoidal to elongated, present in the perianth midveins, ovary and 
capsules. Calyx cupulate, equaling or somewhat longer than corolla tube, rarely 
in some flowers shorter than corolla tube, divided 1/2-2/3 of the length, lobes 
triangular-ovate, acute, not or only slightly overlapping at the base. Corolla tube 
campanulate to suburceolate, 0.5-1.3(-L.5) mm long, lobes 0.8-1.2(-1.5) mm long, 
triangular-ovate, acute, ca. 1/3 to equaling corolla length, suberect to erect, 
apically inflexed. Stamens barely exserted or enclosed; anthers (0.2-)0.3-0.45 
x 0.19-0.25 mm long; filaments equaling or longer than anthers; pollen as in C. 
indecora. Infrastaminal scales ca. reaching the filaments, united with the co- 
rolla tube for most of their length, oblong, bifid, with short dentate wings or 1-3 
fimbria on each side of filament attachment, rarely truncate with 3-6 fimbria. 
Styles distinct, 0.7-1.5 mm long, + unequal, evenly filiform, strongly divergent 
in capsule; stigmas capitate, globose. Capsules dark-brown and not semi-trans- 
parent when dried (pericarp thick), glabrous, initially globose later become 
evidently depressed, 1.6-2.4 times wider than long; the former suture lines be- 
tween the 2 carpels forming a longitudinal groove on opposed sides of capsule; 
depressed and thickened around the style bases, indehiscent or irregularly de- 
hiscent, surrounded or capped by the withered corolla; pericarp epidermis with 


218 BRIT.ORG/SIDA 22(1) 


ahh, “" 


Gi ee 


$2 


Fic. 2. Floral variation in C. coryli: a—g. excepting e (scale bar = 1 mm); e. Convex fleshy epidermis cells in the corolla 
(scale bar = 150 um). 


COSTEA ET Al OF CUSCUTA INDECORA COMPLEX 219 


Fic. 3.C | | 1s of C. indecora and C. coryli.a. Capsule of C. ind (scale k mm); b. Capsule of C. coryli 
(scale bar = = 1mm); « Surface of capsule of C. coryli (scale bar = 70 Lim); fe -e, eae i seeds of C. indecora: d. 
Dorsoventrally compressed; e. More or less angled (scale bar = 0.60 mm); f-g. Surface oat gees f.C. col polyge: 


Dal 1 ith £ 


nal with epicuticular wax; g-h. C. indecora: g. Alveolate; h. Polyg with groups of pap 


220 BRIT.ORG/SIDA 22(1) 


a prominent pattern of polygonal cells. Seeds 3-4 per capsule, 1.32-1.65 x 1.25- 
1.4mm, similar to those of C. indecora; additionally seed coat with polygonal 
epidermal cells may have epicuticular wax as in Fig. 3f. 2n =? 

Distribution and ecology.—“Throughout the United States east of Rocky 
Mountains, but less common southward and westward” (Yuncker 1965). 
CANADA: Manitoba, Ontario, Québec, Saskatchewan. U.S.A.: Alabama, Arkan- 
sas, Arizona, Connecticut, District of Columbia, Delaware, Iowa, Illinois, Indi- 
ana, Kansas, Kentucky, Maryland, Massachusetts, Michigan, Minnesota, Missis- 
sippi, Missouri, Montana, Nebraska, New Jersey, New Mexico, New York, North 
Carolina, North Dakota, Ohio, Oklahoma, Pennsylvania, Rhode Island, South 
Carolina, South Dakota, Tennessee, Texas, Virginia, Wisconsin, West Virginia. 

The floristic record from Québec is new, based on a single collection: 
QUEBEC, Cham bly Co. St. Lambert, 9 Aug 1935, Terrill 884 (MTMG). The spe- 
cies is considered “critically imperiled” (SL) in Canada (Argus & Pryer 1990; 
NatureServe 2004). Flowering Aug-Oct. Hosts: wide range of herbaceous and 
woody species, including Aster, Ceanothus, Corylus, Helianthus, Monarda, Rhus, 
Rubus, Solidago. 

Conservation status.—S2S3 Gmperiled to vulnerable) in the U.S.A. (not yet 

assessed by NatureServe 2004); critically imperiled (NI) in Canada (Argus & 
Pryer 1990; NatureServe 2005). 


— 


3. Cuscuta warneri Yuncker, Brittonia 12:38. 1960 (Fig. 4.). Type: U.S.A. UTAH. Millard 
Co. vicinity of Flowell, 15 mi W of Fillmore, on Phyla cuneiformis, 10 Sep 1957, Warner s.n. 
(HOLOTYPE: UTC; tsorypes:; DAO, DPU, GH, NY, OSC, RSA, US, WSU) 

Stems 0.30-0.50 mm thick, yellow. Inflorescences of subsessile flowers on gla- 

brous pedicels, 0.5-1 mm, in few-flowered glomerules; bracts one at the base of 

pedicels, ovate to lanceolate. Flowers 5-merous, 2.1-4 mm long, slightly fleshy, 
white-creamy, papillate-hispidulous, corolla epidermis with papillae 30-50p1m 
long oriented in rows; epicuticular wax consisting from longitudinally reticu- 
lated rodlets; laticifers isolated or in longitudinal rows as in C. indecora. Calyx 
campanulate-cupulate, ca. 1/2 the corolla length, divided ca 1/2, lobes triangu- 
lar-ovate, carenate, each apically enlarged to form a large, prominent, diver- 
gent, horn-like projection, 0.5-0.75 mm long, and sometimes basally with a 
smaller multicellular projections; not overlapping. Corolla tube campanulate- 
urceolate, 1.7-2.5mm long, lobes triangular-ovate, more or less auriculate, acute, 
0.5-0.7 mm long, 1/3-1/4 the corolla length, suberect, apically inflexed and 
basally overlapping. Stamens included, incurved over the ovary, anthers broadly 
elliptical 0.4- 07 x 03-0. 4mm; filaments about as long as the anthers; pollen 
as in C. indecora. Infrastaminal scales ca. reaching the filaments, united with 
the corolla tube for ca. 1/2 of their length, oblong, shallowly and irregularly 
toothed at the truncate apex. Styles distinct, 0.2-0.4 mm, evenly filiform, barely 
longer than the collar-like stylopodium; stigmas globose, capitate. Capsules 


— 


COSTEA ET AL., TAXONOMY OF CUSCUTA INDECORA COMPLEX 221 


Fic. 4. Cuscuta warneri. a—b. Morpholedy pies a. SANE, bs Sena (scale bars = 1 mm); ¢. Papillae on 
corolla bale Pals 75 um); d. C mon): e-f. Morphology of seeds: e. 


| wal Vek Cork £h Ieatad + va / 
yy y P 5 Lm) 


i, VETTEL GT VIEW (oCdle Val 


222 BRIT.ORG/SIDA 22(1) 


yellowish, + semi-transparent when dried, more or less papillose, globose, 0.9- 

2 wider than long, the suture line between the 2 carpels not depressed; thick- 
ened and raised around the style base, indehiscent or irregularly dehiscent, en- 
veloped by the corolla; pericarp epidermis smooth, with scattered papillae 10-20 
jum long. Seeds 2 per capsule, 1.33-1.56 x 1.26-1.40 mm, dorsoventrally com- 
pressed, subround to broadly-elliptic, hilum subterminal, round 0.15-0.18 mm 
in diameter, vascular scar, 0.04-0.08 mm long, linear, oblique; surface of the 
seed coat alveolate when dry and papillose when wet, seed epidermis cells 0.26- 
0.40 um in diameter. 2n =? 

Distribution and ecology.—U.S.A.: The species has been considered “possi- 
bly extinct” because despite repeated search it has never been found again at 
the type locality in Utah (Reveal & Cronquist 1984; NatureServe 2005). How- 

ever, we have found one more collection from southern New Mexico (Sierra Co: 
Pedro Armendaris Grant, 15.6 mi N of Engle, E of Red Lake, 4800 ft, on Phyla 
incisa, 24 Sep 1998, Peterson 98-699 (NMC)). The species has also been men- 
tioned from Arizona (NatureServe 2005), but we are not aware of any herbarium 
vouchers. Albeit clearly extremely rare and endangered, this species might be 
potentially distributed at a low frequency over a larger geographic range, span- 
ning Utah, Arizona, and New Mexico. Flowering and fruiting Jul-Sep. Hosts: 
known only from Phyla sp. 

Conservation status.—T1 (critically imperiled) (GH, presumed extinct, 
NatureServe 2005). 


ad 


4. sass ae Yuncker, Illinois Biol. Monogr. 6:149. 1921. (Fig. 5). Typr: U.S.A. 

ALIFORNIA: Big Horse Mountain, South Fork of Eel River, 3 Aug 1892, W_L. Jepson 5c (HOLO- 

TYPE: JEPS, fragment NY). 

Stems 0.30-0.40 mm thick, pale-yellow. Inflorescences of short-pedicellate flow- 
ers in cymose clusters; bracts one at the pedicel base, ovate to lanceolate. Flow- 
ers 5-merous, 2-2.7(-3) mm long, fleshy, white-cream; epidermal cells arranged 
in rows, fleshy, with anticlinal walls convex (dome-like), additionally with 
cylindric-conical papillae 40-70 um long; when flowers dry up, rows of dehy- 
drated epidermal cells are usually easily discernible; epicuticular wax repre- 
sented by longitudinally reticulated rodlets; laticifers isolated ovoidal to elon- 
gated, or arranged in longitudinal rows, present in the perianth along the 
midveins, ovary and capsules often running longitudinally and yellowish-or- 
ange colored. Calyx shallowly cupulate, ca. 1/2 as long as the corolla tube, di- 
vided ca. 1/2 the length, lobes triangular, acute, not basally overlapping. Co- 
rolla tube campanulate-globular, becoming suburceolate, 1.3-2 mm long, lobes 
triangular, acute, less than 1/2 as long as the tube, erect, with inflexed apices. 
Stamens mostly included, anthers broadly elliptical 0.2-0.3 x 0.1-0.2 mm long: 
filaments about as long as the anthers; pollen as in C. indecora. Infrastaminal 
scales lacking or represented only by ridges and short bridges. Styles 0.4-0.8 


COSTEA ET Al OF CUSCUTA INDECORA COMPLEX 223 


Fic. 5. Cuscuta jepsonii. a. Flower (scale bar = 1 mm); b. Pollen (scale bar = 10 im). 


mim long, + unequal, somewhat subulate, erect or divergent in capsule; stigmas 
capitate, globose. Capsules light-brown + semi-transparent when dried (peri- 
carp thin), glabrous, subglobose, globose, to slightly depressed-globose, 1-1.5 
wider than long, narrowed and thickened around the style bases, indehiscent 
or irregularly dehiscent, surrounded by the withered corolla. Seeds (no mature 
seeds were seen) 2-4 per capsule. 2n =? 

Distribution and ecology—Known only from the type collection. Flower- 
ing summer-early fall Jul-Sep). Hosts: Ceanothus. 

Conservation status.-GH (presumed extinct) (not mentioned in 
NatureServe 2005). 


APPENDIX 1.—VOUCHERS FOR THE SEM STUDY 
(NY, EXCEPT C. WARNERI FROM GH AND C. JEPSONII FROM UC) 


1a. Cuscuta indecora var. indecora (19 ae pou hae ARIZONA. Pinal Co.: Su- 
pa Gloke Hwy, July 1926, Moore s.n. CA RNIA. Kern Co.: Bakersfield, 400 ft, 6 Jul 1920, Fisher 
. San Bernardino Co.: San Bernardino nts lawer ee of Upper Sonoran Zone, 4000 ft, 9 Nov 
joe Wolf 4392. COLORADO. Larimer Co.: ies Collins, 5000 ft, 25 Aug 1896, Baker s.n. FLORIDA. 
Putnam Co.: 2.5 mi S of San Mateo, 26 Jul 1961, Godfrey & Reinert 61139a. IDAHO. Gooding Co.: 
Hagerman Valley, 13/E, 7S,21 Aug 1941, Davis eee Co.: 2 mi E of Glenns Ferry, 23 Aug 1940, 
Christ 11779. LOUISIANA. Terrebone Parish: around Louisiana Universities Marine Consortium lab 
buildings and along La. 56 in Cocodrie, S of Houma, T215,R18E, 12 Aug 1989, Thomas 111952. MIS- 
SISSIPPI. Harrison Co.: Ship Island, 15 Jun 1952, Demaree 31920. NEBRASKA. Arthur Co.: Arapaho 
Prairie, T18N R39W Sect 31, 32, ca. 1200 m, 27 Jul 1977, Vescio & Kruse 174. NEVADA. Nye Co.: Rt. 52 
near Rt. 16 junction, 2600 ft, 26 Sep 1970, Beatley s.n. NEW MEXICO. Chaves Co.: Bottomless Lakes 
State Park at the edge of ce Lake, 5 Oct 1966, Crutchfield 2319. Eddie Co.: a few m N of Texas 
fe ca.0.5 mi SE of Hw -180, ca. 3900 ft, 1 Sep 1985, Spellenberg & Spurrier s.n. OREGON 
atillaC Peon oon 1944, Peck 22633. TEXAS. Angelina Co.: near Shawnee, 10 Sep 
as ae. ae 11905. El Paso Co.: along Hwy. 62-180, 4 mi E of junction with Hwy 659, ca 


224 BRIT.ORG/SIDA 22(1) 


4000 ft, 23 Oct 1983, Worthington 11583. Llano Co.: hills above Inks Dam, 21 May 1940, Lundell & 
Lundell 9025. UTAH. Utah Co.: 1105, Sec. 4,0.5 mi E of Genola turnoff on Hwy 50-6, 4650 ft, 8 Sep 
1984, Baird et al. 1513. Weber Co.: Howel Experimental Fruit Farm, Pleasant View, N Ogden, 7 Sep 
1967, Nye s.n. 

1b. Cuscuta indecora var. longisepala (2 collections examined).— TEXAS. Chambers Co.: 
Anahuac, 3 ft, 10 Jun 1933, Fisher s.n. Cameron Co.: 4 mi NW of Brownsville, bordering the Military 

wy, 10 m,9 Jul 1941, Runyon 2819 

1c. Cuscuta indecora var. attenuata (3 collections examined)—KANSAS. Republic Co.: 2 mi N 
and 2 miW of Wayne, 13 Sep 1952, Horr 4410. TEXAS. Cameron Co.: Robb's Ranch,0.5 mi N of Ranch 
house, bordering the road, 10 Aug 1941, Runyon 2873, Dallas Co.: Dallas, Sep 187(?4), Reverchon s.n. 


. Cuscuta coryli (17 collections examined)—CANADA. ONTARIO(?): St. Clair River (?), Squirrel 
Island, 16 Sep 1920, Farwell 5692.U.S.A. ARIZONA. (no county given): Grand Canyon, 7 Sep 1886, 
Eggert s.n. ILLINOIS. Menard(?), no date, Hall s.n. INDIANA. Lake Co.: just S of Pine, 19 Sep 1926, 
Deam 43763. Nobble Co.: 4 mi N of Kendallville, 23 Aug 1928, Deam 46128. MARYLAND. 

m o.: wood near Widewaters, 26 Aug 1934, Killip 31293. MICHIGAN. Kalamazoo Co.: 
NE of Schoolcraft, 6 Sep 1938, Hanes 548. MISSOURI. St. Louis, Sep 1842, Engelmann s.n. Bush ae 
Eagle Rock, 28 Sep 1896 and 14 Aug 1905, Bush 202 and 3244. NEBRASKA. Richardson Co.: woods 
5 Lee’s Ranch, 1.5 mi NW of ae i000 ft,15 Sep 1940, Reynolds 2727.NEW JERSEY. Somerset Co.: 

econd Mountain, Watchung, 29 Aug 1937, Moldenke 10086; Little Snalie Hill, Sep 1915, MacKenzie 
NEW YORK. Tioga Co.: Campville, 20 Sep 1895, collector illegible (FE. Fr...) 296; Long Island, Sea 
Cliff, 24 Sep 1928, Ferguson 7181; Staten Island, 24 Oct 1891, Vail s.n. TENNESEE. Carter Co.: Roan 
Mountain Station, 28 Aug 1908, Rydberg &179. WISCONSIN. Madison, no date, Watson s.n. 


3.Cuscuta warneri (1 collection examined),—U.S.A. NEW MEXICO Sierra Co.: Pedro Armendaris 
Grant, 15.6 mi N of Engle, E of Red Lake, 4800 ft, on Phyla incise, 24 Sep 1998. Peterson 98-699 (NMC). 
UTAH. The type collection (GH). 


4. Cuscuta jepsonii (1 collection examined).—U.S.A. CALIFORNIA. The type collection 


UC). 
ACKNOWLEDGMENTS 


We thank directors/curators from ACAD, ALTA, ARIZ, ASU, BRIT, DAO, E GH, 
HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, RSA, 
SASK, 5FS, TEX & LL, TUR UBC, UC & JEPS, UNB, UNM, US, USAS, UWO, 
UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Special thanks to 
NY staff for approving and preparing the four voluminous loans containing 
the herbarium of T.G. Yuncker. Dan Austin, Alan Prather and Lytton Musselman 
provided valuabl t 1 suggestions for an earlier version of the manu- 
script. Special appreciation goes to Therry Deroin (P) for sending the fragments 
of the Cuscuta indecora type and to Barbara Ertter (UC) and Emily Wood and 
Walter Kittredge (GH) for permission for SEM study of the types of C. jepsonii 
and C. warneri. Alexandra Smith assisted us with the scanning electron micro- 
scope. Elma Schweigert translated the abstract into Spanish. 


REFERENCES 
Arcus, G.W. and K.M. Pryer. 1990. Rare vascular plants in Canada: our national heritage. 
Canadian Museum of Nature, Ottawa. 
Be.iz, T. 1986. A revision of Cuscuta sect. Cleistogrammica using phenetic and cladistic 


COSTEA ET AL., TAXONOMY OF CUSCUTA INDECORA COMPLEX 225 


analyses with a comparison of reproductive mechanisms and host preferences in spe- 
cies from California, Mexico, and Central America. Ph.D. diss., Univ. of California, Berkeley. 

Beuiz, T. 2002.Cuscuta.|n:B.G. Baldwin, S. Boyd, B.J.Ertter,R.W. Patterson, T.J.Rosatti, D.H.Wilken., 
and M.Wetherwax (eds.). The Jepson desert manual. Vascular plants of southeastern 
California. Univ. of California Press, Berkeley. Pp: 280-281. 

Costes, M., G.L. Nesom, and S. Steranovic. 2006. Taxonomy of the Cuscuta pentagona com- 
plex (subsect. Arvenses: Convolvulaceae) in North America. Sida 22:151-175. 

ENGELMANN, G. 1842. A monograph of the North American Cuscutineae. Amer. J. Sci. Arts 
43:333-346. 

ENGELMANN, G. 1843.Corrections and additions to the monograph of Cuscutineae. Amer J. 
Sci. Arts 45:74-78. 

ENGELMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with 
critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 
1:453-523 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. First edition. /n: Kartesz, J.T.and 
C.A.Meacham. 1999. Synthesis of the North American Flora, Version 1.0.North Carolina 
Botanical Garden, Chapel Hill, NC. 

NatureServe. 2005. NatureServe Explorer:An online encyclopedia of life [web ep plication 

r> 


K++ 


Version 4.0. NatureServe, Arlington, Virginia. <http:// rg 


(Accessed: November 16, 2005) 

Parker, C.and C.R.RicHes. 1993.Parasitic weeds of the world. Biology and control. CAB Inter- 
national, Wallingford, UK. 

Pazy, B. and U. Puitmann 1995.Chromosome divergence in the genus Cuscuta and its sys- 
tematic implications. Caryologia 48:173-180. 

Pinkava, D.J., R.K. Brown, J.H. Linosay, and L.A. McGit. 1974. |OPB Chromosome Number Re- 
port XLIV. Taxon 23:373-380. 

Prater, L.A.and J. Tvat. 1993. The biology of Cuscuta attenuata Waterfall. Proc. Okla. Acad. 
Sci. 73:7-13. 

PratHer, A.L., RJ. Tyat, and W.D.Warbe. 1995.A taxonomic i tigation of Cuscuta attenuata 
(Cuscutaceae) and related taxa. Sida 16:447-458. 

Revea, J.L. and A. Cronaquist. 1984. Cuscutaceae. In: A. Cronquist, A., A.H. Holmgren, N.H. 
Holmgren, J.L. Reveal, and PK. Holmgren. Intermountain flora, Vol.4. New York Botanic 
Garden, Bronx. Pp. 77-84. 

WarerrAaLt, U.T.1971.New species of Cuscuta and Phlox from Oklahoma. Rhodora 73:575- 
Dis 

Yuncker, 1.G. 1921. Revision of the North American and West Indian species of Cuscuta. 
Illinois Biol. Monogr. 6:91-231.Reprinted 1970, Johnson Reprint Company, N.Y. 

Yuncker, 1.G. 1932. The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. 

YUNCKER, T.G. 1960. Two new species of Cuscuta from North America. Brittonia 12:38-40. 

YUNCKER, T.G. 1965. Cuscuta. North American Flora, ser. 2,4:1-51. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


RICHARD C. BEIDLEMAN. 2006. California’s Frontier Naturalists. ISBN 0520230108, 
hbk.). University of California Press, Berkeley. (Orders: California Princeton 
Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. Tel: 
609-883-1759; Fax 609-883-7413). $39.95, 484 pp., b/w illus. map, 5 1/2" 
x 91/2" 

In this well-written and very readable book, Dr. Beidleman offers the history of California natural 

history exploration through the lives of those men and women who ardently and assiduously ob- 

served and collected throughout the state. They sought and they found the unknown, new species 
never before described. Data of their lives, their backgrounds, and their motives are detailed. Their 
lengthy. 


personalities are revealed by anecdotal passages, some no more than a sentence or two, some 

Coverage begins with exploration around Monterrey in 1786 by the Conte de La Pérouse under 
the French flag and ends in the early twentieth century with the remarkable John and Sarah Lem- 
mon and Kate Brandegee. In between are numerous explorers assigned to the boundary and railway 
surveys and particularly the prodigious California Geological Survey (1842-1874). Some of the sci- 
entiste xplorers were instrumental in tk tablishment of such institutions as the University of Cali- 
ornia, Berkeley, the California ae of Sciences, and the Lick Observatory. 

While references include the whole range of biology, geology, and astronomy, many of these 
frontier naturalists were botany-minded. Their findings included hundreds of new species which 
were sent to Ha Page ae Asa ie ana Jone ae iy cate author names many of these plants, 
plant in its family. However, 


eis 


usually w 
those not versant in California flora might be ae oer to have on hand afield guide if not the Jepson 
manual for such citations as “the Pale- Yellow Layia,” or the “poisonous prickly poppy, Chicolote.” 
_ ris neo one meets ee characters Wiheee names may not be in common parlance 
I f um lemmonti, Carex breweri, Baccharis plummerae, Lilium 


ae Tritonia palmeri among oe 
If Jo ohn Nt seems to have Peet granted eave short a ift, the: author cet tainly recognizes 
hi vation. In general Dr. Beidleman 


feauunes tae naturalists who cone bet d significantly in se enunic discovery.—Joann Karges (re- 
tired TCU Library), Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(1): 226. 2006 


A NEW SPECIES OF PEDIOMELUM (FABACEAE) 
FROM THE LOWER PIEDMONT PLATEAU OF GEORGIA 
AND SOUTH CAROLINA 


James R. Allison Michael Wayne Morris 
ox 511 PO. Box 2583 
Rutledge, Georgia 30663, U.S.A. Gainsville, Georgia 30503, U.S.A. 


oun N. Egan 


AA 


Dept. of | licrobiol nd Mol leculal Biology 
sisham ie University 
Provo, Utah 84602, U.S.A 


ABSTRACT 
In the Piedmont Physiographic Province of the southeastern U.S.A., the endemic North American 
genus Pediomelum is inown a rom three dry, rocky, partly open sites near the Fall Line. The first 
collections were made in 1984, from Richland County, South Carolina; in 1996, from more than 100 
km. to the west, in Si County, Georgia; and in 2005, from Lexington County, South Carolina, 
less than 20 km from the Richland County site. The two late-twentieth-century collections were 


— 


referred, with reservations, to P. canescens, a species of sandy soils on the adjacent Atlantic Coasta 


Plain. This was the only known Pediomelum that resembled the Piedmont plants in having the peti- 


oles shorter than the puoulesanat the only similarly eee cauesecat species east of the Missis- 
sippi River. Sul t 1 study by tl 


that the Piedmont populations 
-onsistent Penola y that is unique within the genus in combining subsessile leaves with 

congested, many-flowered inflorescences. These plants differ from P canescens in additional ways 
(e.g., fruiting calyces gibbous and more narrowly campanulate, bracts conspicuously ae and 
broader, and leaflets more narrowly elliptic). The Piedmont plants also cannot be considered a sessile- 
leaved variant of any of the western species, and therefore they are described as Sane m 
piedmontanum Allison, Morris, & Egan, sp. nov 


share a 


RESUMEN 
En la Provincia Fisiografica de Piedmont del sureste de EEUU, el género Pediomelum, endémico de 
Norteamerica, se ha encontrado solamente en tres sitios secos y rocosos y parcialmente abiertos que 
estan cerca de la Fall Line. Las primeras colecciones se hicieron en 1984 en el Condado de Richland, 
Carolina del Sur; en 1996, a mas de 100 km al oeste, en ae peace ee Comme Georgia; y en 2005 en 
el Condado de Lexington, Carolina del $ “ondado de Richland. Las 


dos colecciones de finales del siglo XX se refirieron, con reservas, a P canescens, una eae de los 
suelos arenosos en el Atlantic Coastal Plain contiguo. Este era el tnico Pedi 


asemejaba a las plantas del Piedmont en tener los peciolos mas cortos que los pecidlulos y la unica 
especie semejante al este del Rio Mississippi erguida y caulescente. Las colecciones subsiguientes y 
los estudios por los autores indican que las poblaciones del Piedmont comparten una morfologia 
coherente que es unica dentro del género en combinar las hojas subsésiles con las inflorescencias 
ene y multi sre Estas plantas se diferencian de P canescens de otros modo p. ej., los calices 

] 


\ | las bractea 


} fe} £ 


ce Fe | 


bi mas 


5 


SIDA 22(1): 227 — 241. 2006 


228 BRIT.ORG/SIDA 22(1) 


anchas, y las hojuelas mas angostamente elipticas). Las plantas del Piedmont tampoco se pueden 


caus wr como una variante, con las hojas subsésiles, de cualquiera de las especies occidentales, y 


mon 


por lo tanto se describen como Pediomelum pier anum Allison, Morris, & Egan, sp. nov. 


HISTORICAL SKETCH 

“This specimen is an enigma.” So began Duane Isely (1918-2000) in his 199] 
annotation of J.S. Angermans.n.(USCH), a South Carolina specimen whose origi- 
nal label bears a date une 8, 1984) but lacks any name for the plant. Above this 
label is an annotation slip with James Grimes’ 199] determination as 
Pediomelum cuspidatum (Pursh) Rydb., made the year after publication of his 
monograph (1990) of Pediomelum Rydb.and other genera of New World legumes 
often treated at the time as components of a single broadly-defined genus, 
Psoralea L. Grimes determination of Angermans.n. as P. cuspidatum must have 
preceded Isely’s annotation, which was manifestly written in dissent: 


~~ 


This specimen is an enigma. It is not Pediomelum cuspidatum, which is a species ol the Great 


Plains The leaves of P cuspidatum are ane and 3- ei liclate These are sess sile and 3-{oliolate. The 


only Pediomelum currently known in South Carolina is P canescens. It matches this to the extent 
that the leaves are shortly petioled or subsessile, but the inflorescence is entirely different 
Clearly as a result of seeing these remarks, Grimes took another look at 
Angerman s.n., making some floral dissections and measurements of the parts 
(as recorded in pencil drawings on the sheet) that were the basis for his second 
annotation, which he attached above Isely’s, with reasons why the specimen 
also did not fit typical Pediomelum canescens (Michx.) Rydb. (pubescence, leat 
shape, number of floral nodes, length of lower calyx-tooth and of wing-petals). 
Before concluding this annotation with the results of his floral dissection, 
Grimes wrote that the plant probably merits varietal rank. We assume he meant 
under P canescens, since he confined his comparison to that species, sensu stricto 
though his unelaborated determination of three months earlier as P cuspidatum 
was not explicitly retracted. 

In 1996, five years after the Isely and Grimes annotations but without 
knowledge of them, Thomas S. Patrick of the Georgia Natural Heritage Program 
(GNHP) made the next known collection of a Pediomelum from the Piedmont 
Plateau (Patrick sn. GA, NY). It came from a locality with unusual geology 
(serpentinite and eels rocks) near the Savannah River in Columbia County, 
Georgia. Patrick identified the legume using Radford et al. (1968), in which, due 
to the palmately foliolate and essentially sessile leaves, it keys to Psoralea 
canescens Michx. (= Pediomelum canescens), a species known from the nearby 
Atlantic Coastal Plain sandhills of Georgia and South Carolina, as indicated in 
Radford et al. That the habitat of Patrick s.n. was a rocky place in the Piedmont 
did not militate against such an identification, as the site supports a natural 
community with a mix of Piedmont and Coastal Plain flora. Among the latter 
are Pinus palustris P. Mill. longleaf pine), very rare in the eastern two-thirds of 


ALLISON ET AL.,A WEY OF EXILS UE 229 


the Piedmont of Georgia, and Marshallia ramosa Beadle @ FE. Boynt. 
(Asteraceae), a state-protected rarity otherwise endemic to the Coastal Plain. 

In June 1999 Morris encountered this same Georgia population of 
Pediomelum and collected Morris 4558 (Herbarium of North Georgia College 
and State University, “NGCSU”). At the time, he was unaware of both the 1984 
specimen at USCH and Patrick’s Pediomelum collection (then stored at GNHP). 
After trying to identify his collection using Isely (1990) and finding that, be- 
cause of the leaf shape and dense inflorescences (Figs. 1, 2), the plant did not 
key straightforwardly to any species, Morris showed his collection to Allison, 
who had the benefit of familiarity with P canescens. Noting at once the much 
showier inflorescences and proportionately narrower leaves of Morris’ collec- 
tion, Allison believed it must either represent a new species or a long-range 
disjunct. After obtaining the two most recent monographs of the group 
(Ockendon 1965; Grimes 1990), Allison, Morris and Patrick agreed that the plants 
appeared to represent a new species. An abstract reporting these findings, but 
without knowledge of the specimen from South Carolina, indicated the inten- 
tion to name the new species Pediomelum georgianum (Allison et al. 2003). 

In 2004 Egan undertook a study of the phylogenetics, biogeography, and 
diversification of North American Psoraleeae (Egan & Crandall 2005). In ad- 
vance of field sampling of the group, she made inquiries about localities with 
recent collections or observations of the various taxa. While inquiring about 
South Carolina localities for Pediomelum canescens, Egan was alerted by the 
curator of USCH, John Nelson, about the “enigmatic” South Carolina collection. 
After borrowing it, she agreed with Isely and Grimes that it was not a clear 
match for any described taxon and sought out the actual living plants. In June 
2005, Egan found the plant at the likely original collection site, in Richland 
County, and also discovered a population about 19 km away in Lexington 
County. The consistent and distinctive morphology confirmed her suspicion 
that it was a new species. Internet research led her toa listing (GNHP 2004) of 
an undescribed Pediomelum on a roster of Georgia rare plants, resulting in her 
contacting Allison. 

After comparison of all the known specimens of Piedmont Pediomelum, 
from Angerman s.n. through Allison’s September 2005 collections of the first 
fruiting material from the South Carolina sites, we have concluded that these 
occurrences of Pediomelum, one in Georgia and two in South Carolina (Fig. 3), 
are all of the same, undescribed species, with a provisional nomen nudum of P 
georgianum. The delay in formal description permits us to give the new species 
a more appropriate specific epithet: 


Pediomelum piedmontanum J.R. Allison, M.W. Morris & A.N. Egan, sp. nov. (Fig. 
2). TyPE: UNITED STATES. GEorGiA. Columbia Co. ca. 12 km NE of Appling. Dixie Mountain, 
20 Jul 2001, James R. Allison & Michael Wayne Morris 12764 (HOLOTYPE: NY; isoTyPes: BRIT, 
BRY, FLAS, GA, GH, MISS, MO, NGCSU, NLU, NY, PH, US, USCH, VSC). 


230 BRIT.ORG/SIDA 22(1) 


Fic. 1. Upper portion of a pl f Pediomelum pied ith the ct | y-flowered inflores- 


I i | ide. Columbia County, Georgia, 15 Jun 3002. 


be | 


Inter spec ies Pediomeli subge sneris Pediomeli sensu Grimesii pe stiolis brevioril p etiolulis ad 


P. canescens solum accedit, autem simul est P. reverchonio sali simile bracteis levalibus magnis et 
latissimis et valde caudatis, sed ab ambobus statim distinguitur inflorescentiis densis et multifloris. 
Plant an erect, strigose and glandular-punctate perennial herb 0.5-0.8(-l) m 
tall; root deep, woody, fusiform, branching, rough-furrowed, to at least 1.5 ¢m 
wide x 15 cm long (Fig. +D). Stems each senescing and detaching at ground 
level promptly after maturation of fruit, including the persistent infructescence; 
in life 1-few, to 6 mm in diameter, branching usually a little below the middle, 
with exfoliating scaly epidermis below, sometimes purplish toward base, stri- 
ate, blond to dark brown punctate-glandular, and strigose (hairs 0.4-0.9 mm 
long), often with remnants of cataphylls, these usually only one or two, remote, 
and some distance from base, 6-L0 mm wide x 6.5-9 mm long, obviously veined, 
sometimes bifid. Stipules erect, persistent, linear-lanceolate, 7-12 mm long x 
6.5-9imm wide, papery, very sparingly strigose, obviously veined, the lower ones 
sometimes fused-free, free above. Leaves palmately 3(-5)-foliolate (Figs. 1, 2, 4B): 
petiole (O-)1-2.4(-4) mm long (reduced upward), terete to more usually slightly 
canaliculate, sparingly strigose, not jointed to the stem and not swollen at the 


— 


ALLISON ET AL., AINDCVW OFENILOS UE 231 


PLANTS OF GEORGIA 
Cotumbia County 


R Alhsan 


Pech , J 
MW Morns, & AN Egan 


Ca 12 km NE of Appling Dry, rocky openings and 
thin woods on Digic Mountain 


HOLOTYPE 


James R Athison & 55544 
M Wayne Moms July 20, 2001 


Fic, 2. Holotype of Pediomelum piedmontanum, prior to its deposit at NY. 


BRIT.ORG/SIDA 22(1) 


232 
on ) coum 
oe ‘Atlanta ' Me ee - [fo | 
el 4 a. “oe 
og / 
} YY 
I po | oe 
. Ew 2» wh 
en! ee nea a 

oe aa ) 1 GERRGIBR™ ze we Atlantic 

) |UEGAUTE Sp 
a 4 7 Ocean 
( ry 
{ J ool 
} f y4 “pe 
BX! 
y 
~~ ir J 
/ t 1 
; | 200 km 


Fic. 3. County outline map showing Georgia, South Carolina, their capital cities, and portions of adjacent states. Coun- 

ties with known occurrences of Pediomelum piedmontanum (all north of but near the Fall Line) are marked with the 

first letter of the sOunty name e (columbia, exington, Mane) aie enalled, Based ona coun: outline mal oF me 
A., apie 


southeastern U.S PI 


from Physical Map of the Southeast, copyright 1967 by Wilbur H. Duncan. 


base, shorter than petiolules or petiole lacking and petiolules attached directly 
to leaf spur; petiolules usually darker in color than petiole, strigillose, L8-3 mm 
long; leaflets narrowly to broadly elliptic, (0.4-)0.6-2.7 em wide X (1.0-)1.2-5(- 
5.5) cm long, apex rounded or shallowly retuse, often mucronate, basally cu- 
iate, upper surface sparingly strigose, especially on midvein, 


= 


neate, strigose-cili 
lower surface usually slightly paler and more abundantly strigose, both sur- 
aces densely dark-glandular. Inflorescence a dense, many flowered 
pseudoraceme, in outline [ovate or] elliptic to oblong, (2.1-)2.5-5.7 cm long, with 
(4-)6-13(-15) nodes and (1-)3(-4) flowers per node; peduncle 6 28Ca 5 cm 
long, base of same color and texture, not jointed to stem; rachis -)2-5(-5.5) cm 
long, elongating only slightly in fruit, usually some or most of the nodes in 
subopposite pairs, internodes (0.5-)1-6(-L0) mm in length; bracts conspicuous, 
enclosing the calyx, persistent, broadly ovate to suborbicular, caudate at apex, 
body 8-11.5 mm wide x (7-)9-10 mm long, densely amber- to dark 


ALLISON ET AL., A NEW SPECIES OF 233 


brown-glandular in age, often purplish-tinged distally, especially the 4.5-7(- 
8) mm long tail; pedicels 1.1-3 mm long, pubescent. Flowers (Fig. 4A) 12.5-14 
mm long; calyx (10-)12-13 mm long to upper teeth, 12-16 mm to lower tooth, 
the tube green or sometimes suffused with violet, 4-5 mm long, nearly glabrous 
internally, externally more or less pilose, at least below the lower tooth, and 
amber- to dark brown-glandular except for the gibbous area, sometimes largely 
eglandular except along the vein running into each of the upper and medial 
teeth, the teeth linear-lanceolate, darker green and often tinged with violet, at 
least distally, copiously pilose-ciliate proximally and sparsely so distally, the 
upper four teeth 4-8 mm long X 1-1.5 mm wide, nearly glabrous on both sur- 
‘aces or internally more or less sericeous, the lower tooth (6-)7-11(-11.5) mm 
long X 15-3 mm wide, internally sericeous, proximally externally pilose, the 
calyx in fruit strongly gibbous-campanulate, with a vein to the apex of each 
tooth and one to each sinus, those to the sinuses bifurcating and continuing as 
submarginal veins more than 3/4 the length of the tooth; petals violet to laven- 
der, or cream to yellowish with tinges of violet on wings and keel; banner obo- 
vate or broadly oblanceolate, (8.5-)10-14 mm long X 5.5-7 mm wide, the claw 
(3.5-)4.5-7 mm long, the blade shallowly emarginate or occasionally truncate, 
low-biauriculate, the auricles scarcely internally callose; wings (7-)8-12 mm 
long X 2-2.5 mm wide, the claw (3-)4-6 mm long, the auricle 1-15 mm long; 
keel petals (5-)6-10 mm long x 2-2.5(-3) mm wide, the claw (3-)4-5 mm long, 
the blade with a darker violet blotch apically, occasionally very slightly apicu- 
late; androecium 9.5-11 mm long, apically calyciform, anthers obovoid-elliptic, 
0.5(-0.75) mm long; gynoecium 8-9 mm long, ovary glabrous, style glabrous or 
sometimes strigillose at the base. Fruit body broadly elliptic to slightly obo- 
vate in profile, 6-7 mm long X 4-4.5 mm wide, glabrous, densely amber- to 
dark brown-glandular, narrowed (but not abruptly) to the arcuate beak, which 
is (5-)6-8 mm long x 2-3 mm wide, glandular like the body, adaxial surface 
sparsely short-pubescent, distinctly longer than the calyx teeth and strongly 
exserted (Fig. 4C). Seed weakly reniform, 3.5-5 x 2.5-3.5(-4) mm, compressed 
but not flattened, nearly smooth, gray-brown (Fig. +E). Flowering late May-late 
June (-late July), fruiting July-August(-September). 

Habitat and range.—Apparently endemic to rocky, open areas and adjacent 
open woodlands in the lower Piedmont Plateau of Georgia and South Carolina 
(Fig. 3). 

Paratypes. U.S.A. GEORGIA. Columbia Co.: (topotypes): 16 Jun 1996, Patrick s.n. (GA, NY); 21 Jun 
1999, Morris 4558 (NGCSU, NLU); 25 Aug 2001, Allison et ee 13001 (GA); 2 Jun 2002, Morris 4971 
(NGCSU, NLU, SWSL); 15 Jun 2002, Allison & Morris 13423 (CLEMS); 5 Sep 2002, Allison 13459 (MO, 
VSC}; 25 Jul 2003, Allison 13611 (FLAS, GH, NY, US), Allison eee: 19 Aug 2005, Morris 5039 
(NGCSU); 24 Aug 200 n is 13777 (NCU); 18 Sep 2005, Williams & Morris 55 (NGCSU); 
27 Sep 2005, Allison 813 (AUA, FSU, GA, JSU, PH, TENN, UNA). SOUTH CAROLINA. Lexington 
Co.: 23 km NW of Columbia, 5 Jun 2005, Egan & Egan 263 (BRY, NY); 25 Sep 2005, Allison 13800 (BRY, 


— 


— 


234 


BRIT.ORG/SIDA 22(1) 


dogs dotsil 


trongly tinged with violet di 
that conceal the calyces except forthe slender cly-teth, th 


‘ | Bem 2002; 
(collected 25 Jul 2003). B:a 5- foliolate lea uncommon in me suonas and unknown in P canescens, but frequent in 
of them ;24A 


ug 2 


h g the 
sera beats if the fits, greener in ig unripe state than the bracts, and longer and broader than the calyx teeth; 
25 Jul 2 


large, arcuate, and strongly 
Allison et al. 13007 (collected 25 Aug 2001). E: seeds; from Allison 13611 
ie 25 Jul 2003). 


ALLISON ET AL.,A NEV SFCUILS VE 235 


CLEMS, MO, TAMU, TEX, USCH, VDB); 26 Sep 2005, Allison 13808 (GH, NY, US). Richland Co.: 26 km 
NW of Lexington, 8 Jun 1984, Angerman s.n. (USCH; labeled, apparently in error, as from Lexington 
County); 5 Jun 2005, Egan & Egan 262 (BRY, K, NY); 25 Sep 2005, Allison 13801 (BRIT, CLEMS, NCU, 
USCH),; 26 Sep 2005, Allison 13803 (BRY). 


COMPARISONS WITH CONGENERS 


Within the papilionoid legumes (whether treated as a family or subfamily), 
two tribes, the Psoraleeae (Benth.) Rydb. and the Amorpheae Boriss., are char- 
acterized by l-seeded, indehiscent fruits and (usually) glandular foliage. The 
Amorpheae produce terminal inflorescences and mostly pinnately divided 
leaves, while the Psoraleeae have inflorescences that technically are axillary 
and leaves mostly palmately divided (Barneby 1977). Taxonomists have treated 
the Psoraleeae as consisting of a single diverse and widespread genus, Psoralea 
(e.g.,Cronquist 1981), or as composed of a narrowly circumscribed Psoralea and 
several genera segregated from it (e.g. Rydberg 1919-1920). In the most recent 
monograph covering North American Psoraleeae, Grimes (1990) took the latter 
approach, following Stirton (1981) in restricting application of Psoralea toa few 
species of the Old World (typified by P. pinnata L.) that have solitary flowers in 
fascicles subtended by lobed cupula, while the inflorescences of the New World 
segregates are pseudoracemes (racemelike in appearance, but some or many of 
the nodes with two or more flowers). Grimes apportioned all of the New World 
Psoraleeae to segregate genera, many of them proposed earlier by Rydberg but 
not as rigorously defined by him. Grimes’ generic reassignments have gained 
broad acceptance, aided by their prompt adoption by Isely (1990). 

The prime characteristic of Pediomelum, according to Grimes (1990), is a 
transverse rupture of the pod (inset, Fig. +A), the base remaining attached to 
the receptacle after the beaked distal portion and seed fall away. The genus was 
divided by Grimes into three subgenera. Leucocraspedon Grimes consists of two 
trailing species of south-central U.S.A. with pinnately-divided leaves, reddish 
to ochroleucous flowers, and a white ridge surrounding the hilum. The other 
two subgenera have mostly palmately divided leaves, flowers mostly lavender 
to bluish-violet, and the hilum not ringed by a white ridge. Subgenus 
Disarticulatum Grimes comprises nine acaulescent or shortly-caulescent spe- 
cies (one composed of two varieties) in which the infructescence disjoints in 
age at the peduncle-base. Only one species of subgenus Disarticulatum occurs 
east of the Mississippi River (Grimes 1990): P subacaule (Torr. @ Gray) Rydb,, of 
calcareous pavement outcrops (cedar glades) of the Interior Low Plateau of Ten- 
nessee and Alabama (Nashville Basin and Highland Rim), and more rarely the 
Ridge and Valley of Alabama (Allison & Stevens 2001) and Georgia (Baskin & 
Quarterman 1970), the Georgia occurrences about 300 km to the northwest of 
the Georgia site for P. piedmontanum. 

The remaining 10 of the 21 Pediomelum species accepted by Grimes, with 


236 BRIT.ORG/SIDA 22(1) 


the addition of P. piedmontanum, constitute subgenus Pediomelum, with per- 
sistent infructescences and acaulescent habit. It is not clear, based on morphol- 
ogy, which of these species is most closely related to P piedmontanum. As the 
morphological data are supplemented with molecular analyses, evolutionary 
relationships within Pediomelum should become clearer. 

Geography and subsessile leaves do not establish a close affinity to the 
nearly sympatric Pediomelum canescens, when weighed against the very dif- 
ferent inflorescences of P. piedmontanum, many-flowered and persistently con- 
gested. The latter are similar, at least in appearance, to those of certain species 
of western U.S.A, where the subgenus is much more widespread and diversi- 
fied. The occasional develoy tin P piedmontanum olf 4- or 5-foliolate leaves 
(Fig. +B) also suggests a closer affinity to some of the western species, as such 
leaves are unknown in P.canescens but typical of such species of the Great Plains 
as P cuspidatum and P. esculentum (Pursh) Rydb. In his monograph, Grimes 
(1990) opined that there are two natural groups within subgenus Pediomelum 
based on calyx morphology: P.argophyllum (Pursh) Grimes, P digitatum (Nutt. 
ex Torr. @ A. Gray) Grimes, P canescens, and P aromaticum (Payson) WA. We- 
ber appear to be a closely related group of species, with broadly and shallowly 
campanulate fruiting calices. Pediomelum piedmontanum, however, witha nar- 
rower, gibbous-campanulate calyx, would fall into the other group from P 


ee 


canescens. 

The following modification of Grimes’ (1990) Pediomelum key to include 
P piedmontanum, requiring only the expansion of a single couplet, provides a 
summary of the most conspicuous distinctions from P. canescens: 


7. Petioles less than 7 mm, or lacking, shorter than the petiolules. 
Inflorescence loose (much of the axis exposed): leaflets 1-3, less than twice as 
oe as wide, petiolules 5-9 mm; of sandy habitats in the Coastal Plain P. canescens 
Inflorescence congested ne axis Usually ee leaflets 3(—5), more than twice as 
long as wide, petiolules 1.8-3 mm; of rocky habitats in the Piedmont P. piedmontanum 
7. Petioles longer than 7 mm, or if rarely o sane always equal to or ae ae 
petiolules datum et al.) 
Demonstrating major differences from Pediomelum canescens ee not test the 


alternative hypothesis that the Piedmont plants constitute a subsessile-leaved 
variant of a more western species. To rule this out, we compared the Piedmont 
plants to the descriptions in Ockendon (1965) and Grimes (1990) of all known 
Pediomelum taxa. We found that there are always multiple characters separat- 
ing the new species from each of its other congeners. For example, P. 
piedmontanum is similar to P. canescens in its strongly beaked fruit, small co- 
rollas, and subsessile leaves; like only P. reverchonii (S. Wats.) Rydb. in its en- 
larged bracts (causing the Piedmont plants to key to that species in Rydberg 
1919-20); and like both P. cuspidatum and P. esculentum in having many-flow- 
ered inflorescences. At the same time, P cuspidatum differs by its much shorter 


ALLISON ET AL., ANEW JPCAICS UP 237 


fruit-beak from both P. esculentum and P piedmontanum, while P esculentum 
has larger corollas, on average, than any of these, besides being uniquely 
eglandular. In short, our comparison of P. piedmontanum with all recognized 
species of Pediomelum indicates that the new taxon is too distinctive to be 
treated at varietal or subspecific rank, under P. canescens or any of the more 
western species. 

Additional research into the systematic relationships, population struc- 
ture, etc. of Pediomelum piedmontanum is underway or is planned for the near 
future, including, as part of ongoing investigations by Egan of the psoraleoid 
legumes, a molecular phylogenetic analysis to determine the closest congener 
for P piedmontanum, anda study of the population genetics of the new species, 
based on DNA sequences. 


HABITAT AND ECOLOGY 


The Georgia and South Carolina localities where Pediomelum piedmontanum 
has been found are all located about 20-25 kilometers north of the Fall Line, 
the name y used for the boundary between the Piedmont and Coastal 
Plain physiographic provinces. At all three of the known occurrences, meta- 
morphic rock is exposed in places, including serpentinite at the Georgia local- 
ity (Cocker 1991la, 1991b) and phyllite at the South Carolina sites (Secor & 
Wagener 1968a, 1968b). Bedrock in these locations appears to be relatively close 
to the surface over an area of at least a few hectares, extending beyond the ac- 
tual outcrops. This is inferred from the plant communities, with the openings 
dominated by drought-tolerant grasses and forbs, such as Schizachyrium 
scoparium (Michx.) Nash and Hypericum gentianoides (L.) B.S.P, and the adja- 
cent, thinly-stocked woodlands dominated by drought-tolerant pines and oaks, 
such as Pinus echinata P. Mill. P taeda L., Quercus falcata Michx., Q. stellata 
Wangenh., and Q. marilandica Muenchh. 

Most of the biomass at all sites is composed of fairly widespread species 
that are frequent in dry, rocky sites of the lower Piedmont. A few relatively in- 
frequent taxa, however, particularly when common to sites in both Georgia and 
South Carolina, may have some value as indicator species (their occurrences 
indicating potential habitat for the Pediomelum). Baptisia bracteata Muhl. ex 
Ell. and Vernonia acaulis (Walt.) Gleason fall into this category. Among species 
recorded from the Richland County site, the uncommon Hypericum lloydii 
(Svens.) P Adams may also have some indicator value. 

The mix of woody plants and particularly the composition of the herba- 
ceous layer suggest that the soil at these Pediomelum sites is somewhat acidic 
and low in fertility. This was borne out by samples of topsoil, each collected 
adjacent to a plant of P piedmontanum, that we sent to the University of Geor- 
gia Soil Testing Lab for analysis, two from the Georgia locale and one from each 
of the South Carolina sites. Because of the small number of samples, we refrain 


238 BRIT.ORG/SIDA 22(1) 


at this time from offering definitive comparisons of soil chemistry among the 
sites, but all samples had an acidic pH value (4.0-6.2). If the new species were 
restricted to the Georgia site, as was formerly believed (Allison et al. 2003), its 
rarity could be easily explained as reflecting adaptation to soil derived from 
serpentinite, a rock rarely exposed in the eastern United States south of Mary- 
land (Tyndall & Hull 1999). The high levels of magnesium and toxic heavy 
metals (e.g, chromium and nickel) characteristic of serpentine barrens were 
reflected in our Georgia test samples, but their concentrations were not nearly 
as elevated in the samples from the two South Carolina sites. For comparison, 
we also submitted a soil sample taken from about 9 km southwest of the type 
locality, from the margin of an outcrop of granite, an abundant rock in the Pied- 
mont. For the pH and the elements named above (Mg, Cr, Ni), as well as for most 
of the other elements measured, the values returned from the South Carolina 
samples were more similar to those from the regionally abundant granite out- 
crop habitat than from the regionally rare serpentinite outcrop of the Georgia 
type locality. In short, the factors responsible for the apparent extreme rarity of 
Pediomelum piedmontanum are not known at present. 

Our observations of the relative vigor of plants indicate that, like the great 
majority of its associates as well as the other members of subgenus Pediomelum, 
P piedmontanum isa heliophyte. Vigor in the new Pediomelum is strongly cor- 
related with the amount of sunlight, with plants growing in the open generally 
larger and more floriferous than the partly shaded plants growing in the adja- 
cent woodlands. The sizeable, deep-reaching root systems of P. piedmontanum 
and its relatives surely provide both a capacity for underground storage that 
aa them against drought, while also placing their shoot-meristems mostly 
out of reach of wildfires, permitting rapid regrowth to take advantage of sud- 
denly reduced competition for light, water, and nutrients. As stated earlier, the 
famously fire-adapted Pinus palustris has persisted at the Georgia type local- 
ity, despite fairly effective exclusion of fire at the site for decades. It seems likely 
that at the South Carolina sites there are (or were at one time) natural openings, 
long-persistent due to shallowness and infertility of soil, that were prevented 
in the past from total closure by occasional fires. It is apparent that the con- 
trolled use of fire should be a primary management tool for the protection and 
enhancement of populations of Pediomelum piedmontanum. 

In terms of pollination ecology, papilionoid flowers like those of 
Pediomelum piedmontanum are considered the product of natural selection 
reflecting entomophily. A detailed study of pollinators has not been undertaken, 
but Morris observed various insects visiting the flowers on June 2, 2002, in- 
cluding bees (probable Bombus spp.) and several species of butterflies, includ- 
ing the pipevine swallowtail (Battus philenor), eastern tailed blue (Everes 
comyntas), buckeye butterfly Gunonia coenia), and dusky winged skippers 


ALLISON ET Al 239 


(Erynnis spp.). The larvae of two of these (Everes comyntas, Erynnis spp.) have 
been recorded as feeding upon one or more legume genera (Klots 1951). 


CONSERVATION STATUS 


Pediomelum piedmontanum appears to be a very rare species. It is difficult to 
imagine that such a striking and distinctive plant, relatively large among the 
herbaceous legumes of its region, could be merely uncommon and yet be rep- 
resented in herbaria by so few specimens. Indeed, publication has been delayed 
until now in part by our concern that the outcome might include its extirpa- 
tion by collectors. That concern has been only slightly reduced by the increase 
in the number of known sites to three, from the single one knownat the time of 
Allison et al. (2003). 

According to NatureServe's global conservation status ranking, a system 
very similar to the IUCN Redlist criteria, Pediomelum piedmontanum would 
be classified as Critically Imperiled (Global Rarity Rank = GI), a rank that 
should be qualified with a question mark (G12), indicating that the taxon is not 
well known enough to assign a rarity rank with confidence. The chief source 
of doubt is the fact that Egan found the Lexington County population quite 
serendipitously, offering hope that there are at least a few other populations 
awaiting discovery. We plan to search for such populations, especially in South 
Carolina. We will also seek partnerships with landowners, land managers, and 
conservation agencies/NGOs to protect and enhance the known localities. 

As there is no guarantee that protection of any of the sites will be possible, 
we have taken the precaution of twice collecting seeds from the type locality 
and providing them to the Atlanta Botanical Garden. The first batch of seeds 
has already yielded a few plants. We intend to work with the ABG and others to 
use these and future production to establish one or more experimental popula- 
tions, in locations near the type locality and with similar edaphic characteris- 
tics. We do not consider that such efforts can be assumed to be sufficient to 
guarantee the long term survival of Pediomelum piedmontanum, however, and 
we urge an expedited consideration of its listing under the federal Endangered 


Species Act and cor responding state laws. 


ACKNOWLEDGMENTS 


Allison is grateful to the Georgia Department of Natural Resources (GaDNR), 
Natural Heritage Program (GNHP), for various forms of assistance, including 
the use of a dissecting scope for the development of the morphological descrip- 
tion; to the Atlanta Botanical Garden for their successful efforts at propagation: 
to the curators and other personnel at GA, GH, FLAS, MO, NY, PH, US, and VSC 
for facilitating the study of Pediomelum collections there; and to José Tallet for 
corrections to the Spanish abstract and to Richard LaFleur of the Department 


240 BRIT.ORG/SIDA 22(1) 


of Classics, University of Georgia, for improvements to the Latin diagnosis. Al- 
lison and Morris are thankful to Thomas S. Patrick of the GNHP for sharing his 
recollections, specimens, and advice, and for companionship in the field. Mor- 
risisalso grateful to the US. Fish and Wildlife Service and the GaDNR for fund- 
ing of the Scutellaria ocmulgee survey during which he first encountered the 
future Pediomelum piedmontanum. He also thanks the Georgia Native Plant 
Society for providing a grant to NGCSU to support an ongoing floristic study 
of the type locality and environs, and he appreciates the dedication and inter- 
est of undergraduates and faculty in the Department of Biology at NGCSU who 
are involved in these and other studies related to the new species. Egan thanks 
John Nelson of USCH for alerting her to the problematic Angerman collection, 
and fora loan that included it and material of P. canescens, very useful for com- 
parison with the new species, and to the curators of AUA, FLAS, FSU, FTBG, 
GEO, MISS, NCU, TROY, USCH, USE VSC, and WFU for providing P. canescens 
locality data. She would also like to thank her advisor, Keith A. Crandall, for 
helping fund her fieldwork, as well as the Society for Systematic Biology and 
the American Society of Plant Taxonomy for support in the form of graduate 
student fellowships. Egan is also grateful to her mother, LaDean Egan, for her 
steadfast assistance and companionship in the field. Finally, all three authors 
wish to thank two reviewers for suggesting valuable improvements to the manu- 
script, Richard LeBlond of the North Carolina Natural Heritage Program, and 
Gregory T. Chandler of the University of North Carolina at Wilmington. 


REFERENCES 


ALLISON, J.R., M.W. Morais, and T.S. Parrick. 2003.Pediomelum georgianum,a new species from 
the Piedmont of Georgia (abstract). Southeastern Biol. 50:185, 

Atwison, J.R.and T.E. Stevens. 2001.Vascular flora of Ketona Dolomite outcrops in Bibb County, 
Alabama. Castanea 66:154—205. 

Barnesy, R.C. 1977. Daleae imagines. Mem. New York Bot. Gard. 27:1-891. 

Baskin, JM. and E. Quarterman. 1970. Autecological studies of Psoralea subacaulis. Amer. 
Midl. Naturalist 84:376-397, 

Cocker, M.D. 1991a. Economic geology of altered serpentinites in the Burks Mountain 
complex, Columbia County, Georgia. Georgia Geol. Surv. Bull. 123. 

Cocker, M.D. 1991b.Geology and geochemistry of altered serpentinites in the Burks Moun- 
tain complex, Columbia County, Georgia. Georgia Geol. Surv. Bull. 124. 

Cronauist, A. 1981. An integrated system of classification of flowering plants. Columbia 
University Press, New York. 

Ecan, A.N. and K.A. Cranpait. 2005. Phylogenetics and biogeography of tribe Psoraleeae 
(Leguminosae) based on nuclear and chloroplast DNA Markers (abstract for 
Systematics Section of ASPT meeting, Austin, Texas). Accessed at www.2005. 
botanyconference.org 


ALLISON ET AL 


241 


GNHP (Georcia Natural Heritace Procram). 2004. Special concern plant species in Georgia. 
Accessed at http://georgiawildlife.dnr.state.ga.us/content/specialconcernplants asp 

Grimes, J. W. 1990. A revision of the New World species of Psoraleeae (Leguminosae: 
Papilionoideae). Mem. New York Bot. Gard. 61:1-114 

Isety, D. 1990.Vascular flora of the southeastern United States, 3(2), Le guminosae (Fabaceae). 
The University of North Carolina Press, Chapel Hill. 

Kiots, A.B.1951.A field guide to the butterflies of North America, east of the Great Plains. 
Houghton Mifflin, Boston. 

Ockenpon, D.J.1965.A taxonomic study of Psoralea subgenus Pediomelum (Leguminosae). 
Southw. Naturalist 10:81-124. 

Raprorp, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill, N.C. 

RypserG, PA. 1919-1920. Fabaceae: Psoraleae. N. Amer. Fl. 24:1-136. 

Secor, D.T.and H.D. Wacener. 1968a. Stratigraphy, structure and petrology of the Piedmont 
in central South Carolina - Carolina Geological Society Field Trip Guidebook 8. (Geo- 
logic Notes, Vol. 12 No. 4). Division of Geology, S.C. State Development Board. 18pp. 

Secor, D.T.and H.D. Wacener. 1968b. Map of Piedmont geology in Central South Carolina. 
Carolina Peciedical Society, Columbia, S.C. (downloaded from http:// 
e icalsociety.org cgscdguide.htm) 

STIRTON, C. H1 1981, sues in the Leguminosae-Papilionoideae of southern Africa. Bothalia 
13:317-25, 

Tynpatt, RW. and J.C. Hutt. 1999, Vegetation, flora, and plant physiological ecology of ser- 
pentine barrens of eastern North America. In:R.C. Anderson, J.S. Fralish, and J.M. Baskin, 
eds. Savannas, barrens, and rock outcrop plant communities of North America. Cam- 
bridge Univ. Press. Pp. 67-82. 


<—s 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


PETR SKLENAR, JAMES L.LUTEYN, CARMEN ULLOA ULLOA, PETER J. JORGENSEN, and MIcHAEFL 
O. DILLON. 2005. Flora Genérica de los Paramos: Guia Ilustrada de las Plantas 
Vascalares. (ISSN 0077-8931; ISBN 089527-468-2). Memoirs of the New York 
Botanical Garden, volume 92. New York Botanical Garden Press, 200" Street 
& Kazimiroff Blvd., Bronx, New York, 10458-5126, U.S.A. (Orders: 
nybgpress@nybg.org). $85.00, 499 pp., 61/2" x 91/2". 


The term eee is io ten misunderstood even in some biological contexts. In this book’s introduc- 
arify the word. Paramos is not an area confined to 


> 


tion, the auth 
northern one “Amen but aie it isan ecosystem a extends from the higher mountains of 


thern Peru, thus embracing these countries as well as Colombia, Ecua- 


ou 


Costa Rica anc 
dor, and Venezuela. It is an ecosystem rich in biodiversity and endemism and one of ecological ex- 
tremes, to which the flora has adapted. Plant characteristics include acaulescent rosettes, shrubs 
with minute leaves, prostrate dwarf shrubs, and geophytes. The purpose of the book, however, is not 
to introduce these plants but to serve students a professionals as a guide to the g of vascular 
plants, ultimately as an incentive for the protection of this unique anc luabl 5 

With this objective in mind, the authors provide many keys: a basic one to each of e on 
owering plants, with especial attention to leaf structures, which ¢ 


groups from pteridophytes to [1 
count for five of the eight general keys. Within each of the 12 families esa aanees eee are 


keys to the 540 genera and keys to tribes. The text ful ly eae ibes the genus and 
and altitudes at which the plants grow. Accompa he text is a picture, in most cases a crawls 
e species in the genus or a repre eseniative aes almost all of these were previously pub- 


— 


of the sing 
lished and are so acknowledged in an appendix. References follow each entr 

The five authors are at eee meena institutions. The leading ae Petr Sklenaf is associ- 
aches at Charles University, Prague. Nine other pro- 
fessionals are cited a rie ee TCU Library), Botanical Research Insti- 
tute of Texas, Fort ate TX 76102- 4060, US. 


:ditor’s note.—See page 724. 


ated with the New York B 


4 


SIDA 22(1): 242. 2006 


BIG-FRUITED BUCKTHORN, SIDEROXYLON MACROCARPUM 
(SAPOTACEAE), A LONG-FORGOTTEN GEORGIA ENDEMIC 


James R. Allison 
PO. Box 511 
edge, Georgia 30663-0511, U.S.A. 
jallison@mindspring.com 


ae 


Rut 


ABSTRACT 
Bumelia macrocarpa Nutt. languished in obscurity for more than 150 years. Thomas Nuttall (1786- 
1859) collected uae low shrub in 1830 and described it in 1849. Asa Gray relegated it to the syn 
onymy of B. lanuginosa (Michx.) Pers. in 1886, apparently due to the lack of subsequent collections 


and without seeing any material of Nuttall’s plant. In 1940 Robert Clark identified type material of 


B. macrocarpa as B. reclinata (Michx.) Vent, sete to notice the fact that more than one species had 


been mounted on the sheet, with only ot all fragment actually representing B. macrocarpa. Since 


Bumelia Swartz is now considered to asynonym of Sideroxylon me the restoration of this distinc- 
tive species, endemic to southeastern Georgia (U.S.A.), requires a new combination, S. macrocarpum 
(Nutt.) J.R. Allison. 


RESUMEN 


Bumelia macrocarpa Nutt. languidecio en la oscuridad durante mas de 150 anos. Thomas Nuttall 
(1786-1859) coleccion6 este arbusto bajo en 1830 y lo describid en 1849. Asa Gray - aesee ala 
1 | 


sinonimia de B. lanuginosa (Michx.) Pers. en 1886, aparentemente debido 
subsiguientes y sin ver ningtin material de la planta de Nuttall. En oe Robert Clark identifico el 
material tipo de B. macrocarpa como B. reclinata cas Vent., al no notar el hecho que se nab 


montado en la lamina mas de una especie, con 
B. macrocarpa. Puesto que Bumelia Swartz s 
la restauracion de esta especie distintiv: lémica al UU), requiere una nueva 


sidera actu: sienenis un sinonimo de Sideroxylon L., 


combinacion, S. macrocarpum (Nutt.) J.R. Allison. 
INTRODUCTION 


Among the genera of woody plants listed in the work that launched the mod- 
ern system of binomial nomenclature, Linnaeus’ Species Plantarum of 1753)! 
was Sideroxylon L. (<Gk. sideros, iron + xylon, wood). In the second edition, of 
1762, he named the first Sideroxylon from the southeastern United States, S. 
lycioides L., and in 1767 a second one, S. tenax L. The species of the temperate 
southeastern U.S.A. (one species extending sparingly to Arizona), spiny shrubs 
or small trees with short styles and finely reticulate, often fascicled leaves, were 
usually treated under the genus Bumelia Swartz for two centuries following the 
latter’s segregation from Sideroxylon in 1788 (e.g. de Candolle 1844; Gray 1886; 
Small 1933; Wood & Channell 1960; Godfrey 1988). An early exception was 


— 


om re tural citat ,see Pennington 


SIDA 22(1): 243 — 264. 2006 


990. 


244 BRIT.ORG/SIDA 22(1) 


Michaux’s (posthumous) description of two species from Georgia in 1803, S. 
lanuginosum Michx. and S. reclinatum Michx. In the following pages, I use one 
or the other generic name interchangeably as deemed appropriate, mostly us- 
ing Bumelia when referring to works which used it, Sideroxylon when discuss- 
ing more recent treatments or concepts. Whatever the botanical name, “buck- 
thorn” has been in use as a vernacular name for these plants for many years 
(Gray 1878). 

All four of these species were restored to their original nomenclature when 
TD. Pennington (1990, 1991) synonymized Bumelia under Sideroxylon as not 
constituting a natural (monophyletic) group. For his coverage of the hardy taxa 
of the southeastern U.S.A., Pennington (1990) was explicit that he relied almost 
exclusively on Arthur Cronquist’s (1945, 1949) publications on Bumelia. Al- 
though at least two dozen species had been named from the USS. in the 150 years 
after Michaux, Cronquist (1945) opined that “not a single valid species of 
Bumelid seems to have been described from the United States since the appear- 
ance of Michaux’s flora in 1803.” Four years after this remark, however, Cronquist 
(1949) described B. thornei Crongq., from collections he had seen from south- 
western Georgia. Relying on Cronquist’s conservative treatment simplified 
Pennington’s (1990) task, as this meant only five temperate species of 
Sideroxylonto be treated, the two each of Linnaeus and Michaux, and S. thornei 
(Cronq.) Pennington? 

However, another monograph covering these species (except Sideroxylon 
thornei), by Robert Clark (1942), had been published just three years before 
Cronquist’s. | consider these almost polar opposites, Cronquist’s a “lumper's” 
treatment and Clark’s that of a “splitter.” For Pennington to have weighed the 
validity of all the taxa recognized by Clark was clearly beyond the intended 
scope of Pennington’s work, which was focused on the Neotropical members of 


—— 


Sideroxylon and other genera of Sapotaceae. 

n Clark’s revision of the U.S. species of Bumelia, he recognized 14 species 
as well as 2 varieties each of B. lanuginosa (Michx.) Pers. and B. lycioides (L.) 
Pers., not including their type varieties. He accepted B. texana Buckland B. rufa 
Raf., raised B. lanuginosa var. anomala Sarg. and var. rigida A. Gray to species 
rank, and accepted most of the species named by J.K. Small. Despite Clark’s 
splitter’s perspective, he did, without comment, reduce to synonymy under B. 
reclinata (Michx.) Vent. one taxon that had been accepted by Asa Gray in the 


2A more distantly related, non-hardy “Bumelia” of the Caribbean has long been known from the warmer areas of 


deen treated everal names, e.g.B 


the southeastern U.S.A, in southern Texas and peninsular Florida. |t has 


cuneata Swartz (misapplied by Gray), B. angustifolia Nutt. (nom. superfl, published 24 years after the next), B. 


celastrina Kunth, and now Sideroxylon celastrint um (Kunth) T.D. Pennington. The still more distantly related 5S. 
salicifolium (L.) Lam. and S. foetidissimum Jacq. are also known from peninsular Florida; they were treated by 
Cronquist (1945, 1946) as Dipholis salicifolia (L.) A.DC. and Mastichodendron foetidissimum (Jacq.) Lam., respec- 
tively. 


ALLISON A 245 


lirst edition of the Synoptical Flora (1878): B. lanuginosa var. macrocarpa (Nutt.) 
A. Gray. Clark’s decision was no doubt based on his examination of the only 
specimen (Fig. |) known to exist, at least in North America, that was identified 
as B. lanuginosa var. macrocarpa or as its basionym, B. macrocarpa Nutt. The 
specimen shows no sign of having been examined by Cronquist, who, like Clark, 
listed B. macrocarpa in the synonymy of B. reclinata. 

[have concluded that Clark’s annotation of the sheet in question as Bumelia 
reclinata resulted from an insufficiently thorough examination of it. The sheet 
is in fact a mixture, bearing four fragments correctly identified by Clark as the 
glabrate B. reclinata, anda single fragment with smaller, thinly but persistently 
strigose leaves corresponding to the protologue (Nuttall 1849) of B. macrocarpa. 
This fragment also matches plants known today from the same region and 
sandy, upland habitat indicated in Nuttall’s protologue. 

In this paper I restore this taxon, a Georgia endemic (Fig. 2), to species sta- 
tus. Although the specific epithet macrocarpa has been used within other 
Sapotaceous genera, it does not appear to be preoccupied in Sideroxylon. There- 
fore the plant's transfer to Sideroxylon requires only a new combination. As this 
taxon has never received a detailed description, I provide one, constructed to 
parallel those in Pennington 1990. 


TAXONOMIC TREATMENT AND DESCRIPTION 


Sideroxylon macrocarpum (Nutt.) J.R. ao cole nov. (Figs. 3-5). BASIONYM: 
Bumelia macrocar pa Nutt. Sylva 3:34. 1849. Bume ur. Madcrocarpa Ae Gray. Sy n. 
Fl. 2(1): 68. 1878. TYPE: U.S.A. GEORGIA (according to the eee “sandy flee not far from 
the Altamaha”): Nuttall s.n. (LECTOTYPE [Fig 1]: PH, accession no. 1030290, the small fragment 
at top left only, which pierces Nuttall’s original label). 


Plants soboliferous ounce less than 1 m high, usually under 0.5 m; geoxylic 
(woody und onger than the aerial, to at least 1O mm thick; 
aerial (leafy) stems B22) mm thick, often with thornlike branchlets bear- 
ing reduced leaves. Long shoots (elongation or leader) tan or reddish-brown and 
tomentulose when young, the hairs blondish or some of them ferruginous, soon 
glabrate, gray with age, armed at most nodes with slender, slightly curved, sharp 
thorns (2-)2.5-13(-20) mm long; some thorns becoming short spur-shoots. 
Leaves tardily deciduous, coriaceous, spirally alternate and spaced at first, soon 
becoming fascicled on short shoots, internodes (1-)2-12(-21) mm long (imper- 
ceptible on short [spur-] shoots); petioles (0.5-)1-3(-5) mm long, channeled, 
pilosulous; blades dark green above, paler beneath, narrowly oblanceolate to 
obovate or spatulate, or broadly elliptic to suborbicular, especially on some spur- 
shoots, (0.3-)1-4(-5.2) cm long, (0.2-)0.4-L5(-2.1) cm wide, glabrate above, per- 
sistently and rather sparsely strigose beneath, occasionally virtually glabrate, 
the hairs blond or rarely slightly rufous; apex obtuse or rounded, sometimes 
retuse, base cuneate or acute; venation brochiodromous or eucamptodromous, 


246 BRIT.ORG/SIDA 22(1) 


Y AG F A — ! Mis 
Paap" P 


Fe| 3 ACADEMY @F SATCRAL ne PUILADELPALA, 
i Bumbo recline, (mich. Vewk Pritsdoiphia 8 das nergendsa Teer, 
pcemmaren or RetukB. Cla. FEB ® 1940 Be ain ty. ee 
as 1, ale only specimen found at PH of Bumeli pa, labeled as Bumelia | va carpa. Inset, 
ge blank sr the sheet: the only material of genuine B | ie Seon maroc 
on the sheet, i t ist I lent | f tl i | 


beginning to expand. The four other fragments are S. jecinatn s. str. 


without a marginal vein, midrib flat or slightly sunken on the upper surface, 
secondary veins mostly 5-8 pairs, ascending, arcuate, intersecondaries usually 
moderately long: tertiaries reticulate-areolate on both surfaces. Flowers bisexual, 
actinomorphic, in axillary (1-)2-15(-18)-flowered fascicles on growth of the 
preceding year. Pedicels 1-3(-5.5) mm long, pubescent. Sepals (4-)5(-6), unequal 
(outer pair slightly s 


— 


norter) (2.1-)2.5-3 mm long, suborbicular to ovate, sparsely 
to moderately pale sericeous-tomentulose, margins scarious, apex rounded to 
subtruncate or slightly retuse. Corolla (4)5(6)-merous, creamy white; tube (0.7-) 


ALLISON A 247 


1-13 mm long, lobes (1.8-)2.4-3(-3.5) % 

mm long, erose; medial lobe-segment os bas, 
erect in anthesis, ovate to suborbicu- Sec eRGuNA 
lar, apex rounded, lateral segments \ | 
(appendages) infolded in anthesis, gag rly = | 


obtuse or sometimes acute, (1.5-)1.8- 
2.3(-3.1) mm long, lanceolate to 
nearly oblong. Stamens (4-)5(-6), 
filaments 1-2 mm long, anthers L.1- 

1.3(-1.5) mm _ long, lanceolate. 
Staminodes (4-)5(-6), infolded, (1.2-) 
1.3-1.8(-2.0) mm long, reaching (1/3) 


. 
4 ‘ 
Waycross 3% rif 


1/2-2/3 the length of the corolla ie ANTI 


midlobes and exceeded by the lateral ee 
wo 

lobes, lanceolate to ovate, apex acute PO 

to rounded. Ovary broadly ellipsoid,  ————— 


f 20 0 2 Miles 


glabrous or medially villosulous. Style ; oe. 
1.0-1.3(-L5) mm long after anthesis, 
glabrous; style-head simple. Fruit 
smooth, black, somewhat lustrous, 
ellipsoid to subglobose, often tipped gp pp 
by the persistent style-base, when Formation, as mapped in Huddlestun 1988. 
fresh 9-12(-14) mm long, 8.5-10 mm 
wide, apex and base rounded, pericarp ca. 2 mm thick, fleshy, sweet. Seed soli- 
tary, (7.3-)8-9 mm long, 5.5-7 mm wide, ellipsoid or obovoid, with truncate base. 
Testa hard, smooth, evenly brown, moderately lustrous, 0.5-1 mm thick; scar 
basal, in two parts, a roughly semicircular area 2-3 mm across and a smaller, 
deltate to lunate abaxial area, the two sometimes joining. 

Phenology and habitat.—Flowering late May through June, or sporadically 
through September. Sandy, well-drained, partly open, pine-oak woodlands and 
pine-oak scrub, southeastern Georgia, U.S.A 


Fic. 2. County outline map of southeastern Georgia show- 
+3 L 1 s42 foe | {A the 


Altamaha River, and three cities. The shaded area corre- 
j L - re "a4 7% + afeok Ale kL 


\dditional coll |: UNITED STATES. GEORGIA: Appling Co.: Co. Rd. 537, 20 Sep 1993, 
Nordman & Tassins.n.(GA); Co. Rd. 308, 20 Sep 1993, Nordman & Tassin s.n. (GH); Co. Rd. 363, 20 Sep 
1993, Nordman & Tassin s.n. (NY); 25 Jul 1996, Allison & Tassin 9352 (FSU, NCU); just SE of Hatch 
nuclear power plant, 26 May 1999, Nelson & Kennemore 20559 (USCH). Candler Co.: GA 40, 27 Sep 
1994, Allison 8606 (USCH); 29 May 1996, Allison 9203 (FSU, GA), 10 Sep 2005, Allison 13782 (CLEMS, 
VSC). Emanuel Co.: 25 mi S of Swainsboro, 24 Apr 1937, Henry 1019 (NY); GA 46, 29 May 1996, Alli- 
son 9200 SU); US 1, 10 Jul 1995, Allison 8914 (FLAS, UNA, USCH); 29 May 1996, Allison 9202 (FSU); 
16 Jun 1999, Allison 12022 (GA, GH, MICH, TAMU, TENN, TROY). Evans Co.: 1.8 mi NW of Bellville, 
7 Apr 1997, Van de Genachte et al. s.n. (PH); 16 Sep 2001, Allison 13043 (GA, GH, NCU, NY, US); 11 Sep 
2005, Allison 13796 (GA, VDB); 1.6 mi NW of Bellville, Allison 9764 (FSU, MO, USCH). Jeff Davis Co.: 
Bullard Creek WMA, 8 May 1997, Allison 9768 (FSU), 8 Jun 1999, Allison 11988 (AUA, BRIT, CLEMS, 
FLAS, GA, GH, JSU, MICH, MO, NCU, NY, PH, TAMU, TENN, TEX, TROY, UNA, US, VDB, VSC); 29 
Sep 1999, Allison 12220 (GA): near Perry Miller Rd., immediately W of Appling Co. line, 26 May 1999, 


248 BRIT.ORG/SIDA 22(1) 


15cm 


RiGee Aeron Oy magocaneti specimen-collegest llecti t GA, all to same scale. A. Two (leafy) aerial st 


lic stem, Allison 11988 B. Part of st with larger than normal 
issues (to5. x <2, 1 cm), Alison 8607, _ Flowering stem, Alison 12040. D. Stem f I g Allison 
lof S$. rufohirtum, Allison 13043 F.S ith tk like t hl 


yt 


bearing reduced leaves, Allison 12022. 


ALLISON 


Fic.4. Sideroxyl nledicect feoedey Te ees 3s eas: oe Tr 
millimeters). A—D, fi hyd I fl f$ (Allison 9203). A. Fl On the right side of the corolla 
tee lpantite asin ofa corolla sabe is evIneNE Bs Sepals, t the outer t I slightl Iler (same scale as 
A). a left corolla “lobe appendage, 2 corlla-lobe and stamen, anda 
right corolla-lobe appendage. D Above pread coroll anther, s n , C= corolla- lobe, lor 
r=left/right append f corolla-lobe ( f the latt letely ol 1 by . Below, upper portion 
PREC TT aan eer 7 are TE og | Alftean 720 


9918). 


§ F & G, comparative material of 
5. rufohirtum: F. Variegated on Meee 1280. G. oral dissection: ob proportionately algae staminades mostly 


250 BRIT.ORG/SIDA 22(1) 


Niele 


Nelson & Kennemore 20576 (USCH). Laurens Co.: GA 19, 21 Aug 1990, Allison 9404 (AUA, BRIT, FSU, 

US, USCH), 18 Jun 1999, Allison 12040(GA, MO). Long Co.: Ft. Stewart, N of GA 144, near Tattnall C 

25 Aug 1992, Zebryk et al. 598 (GA); 8 May 1997, Allison 9766 (USCH, VSC); 9 Jun 1997, Allison a 
(FSU, GA, NY). Montgomery Co.: Co. Rd. 171, 21 Aug 1996, Allison 9408 (USCH); US 280, 21 Aug 1996, 
Allison 9409 (FSU, TEX); GA 135, S of Alston, 22 Aug 19996, Allison 9411 (GA); GA 135, 31/2 mi N of 
Alston, 7 May 1997, Allison 9746 (CLEMS, JSU, PH). Pierce Co.: N of Satilla R., near US 82, 6 Jul 1959, 
Cypert 211 (GA); 2 mi N of Blackshear, 9 Nov 1994, Allison 8754 (FSU); 29 Sep 1999, ae 12228 
(FLAS, GA, TENN, USCH). Tattnall Co.: near Reidsville, 24 Jun 1903, Harper 1851 (GH, 2 sheets; MO; 
NY): NW of Reidsville, 27 May 1957, Broughton s.n. (GA); Eof Ohoopee R., near GA 202, 29 Aug 1985, 
Rayner & McCartney 2401 =e 1); 1987, McCartney s.n. (GA); Gordonia Alatamaha State Park, 


20 Sep 1993, Nordman etal. s.n. (US), GA 57/121 near N limits of Reidsville, 17 Oct 1993, Nordman et 
dl.s.n. (MICH); 27 oe . ie oe (GA); 30 May 1996, Allison 9207 (FLAS, FSU, TAMU, TROY). 
Toombs Co.: 3.8 mi N40'W of Lyons, 24 Oct 1952, Duncan & Hardin 14622 (GA); Co. Rd. 364 at Co. Rd. 


279, 25 Aug 1993, en s.n. (BRIT); 0.2 mi SW of Old Smyrna Cem., 25 Aug 1993, Nordman & 
Tassin s.n. (MO); GA 86 near Pendleton Creek, 17 Sep 1993, Nordman et al. (PH); U.S. 280, 12 Sep 1995, 
Allison 8978 (USCH, VSC); GA 15, 22 Aug 1996, Allison 9418 (GA). Treutlen Co.: Co. Rd. 167, 5 Oct 
1994, Allison 8662 (BRIT, JSU, MO, NCU, NY); 29 May 1996, Allison 9199 (FSU, GA, VDB); GA 86, 23 
Aug 1996, Allison 9430 (USCH). Wheeler Co.: GA 46, 4 Jul 1996, Snow s.n. (GA); 26 Jul 1996, Allison & 
Snow 9356 (GA): GA 19, 21 Aug 1996, Allison 9406 (AUA, FSU, TEX, US, USCH); 17 Sep 2001, Allison 
13048 (GA, MICH, UNA). 


HISTORY OF TAXON 
Nuttall’s choice of epithet, macrocarpa, “large-fruited,” was surely intended to 
contrast with the small fruit of the only other relatively low and small-leaved 
species known to him from the U.S.A., Bumelia reclinata. His treatment of what 
I call Sideroxylon macrocarpum is a fairly accurate, if incomplete, description 
of the plant known today from many more observations, made throughout the 
growing season. The protologue is brief enough to be quoted here in its entirety: 
LARGE-FRUITED BUMELIA. 


BUMELIA MACROCARPA, es ramis Seed valde es spinis elongatis tenuibus 


subrecurvis, foliis parvulis cunea : s obtusis junioribus lanuginosis, demum 
subglabris meu Gus dr upa maxime ovali. 

This very low bushy species, allied to B. reclinata, | give (though from 
very imperfect specimens) to complete the history of our species of 
the genus. The twigs are very slender, at first Se caus covered with 
a grey bark, and with the spines long g and slender as needles. The leaves, 
before expansion, are exceedingly lanuginous, and always small, with 
very short petioles, at length nearly smooth. The fruitis edible, and as 
large as a small date! I found this species on the sandy hills not far 
from the Altamaha, in Georgia, in winter, and therefore do not know 
the flower. It does not grow more than a foot high, and the leaves are 
little more than half an inch long. 


Nuttall’s collection was probably made in 1830. According to Graustein 
(1967), it is only in that year that Nuttall visited the area of Georgia where the 
plant is known to grow today. She also indicates that he was in the Florida Pan- 


+}@n 


ALLISON A 251 


Fic. 5. Sideroxylon macrocarpum in life. Above: unusually extensive patch (10 m? or more); source of Allison 9764, in 
sandy powerline right-of-way, Evans Co., 8 May 1997; note young Pinus palustris, top center. Below locality and 


t 
. L fe Inf £ \ 4 Tl 


? 
acrocarpum 


NO 
c 


dy, w k (j pofS.m 
(indicated by arrow). Inset: branch with leaves, thorns, and a mature fruit, prior to collection as part of Allison 12228, 
Pierce Co., 29 Sep 1999, 


be J 


252 BRIT.ORG/SIDA 22(1) 


handle early in March of 1830, subsequently crossed the Altamaha River|!] et 
route to Savannah, where he departed the Southeast by ship, and that he was 
back in Pennsylvania by April 17. As Nuttall indicated ae he had seen the 
buckthorn only in winter, itis likely that the collection was made in mid-Marc 
When Gray (1878) reduced Bumelia macrocarpa to B. lanuginosa var. 
macrocarpa, not quite three decades after Nuttall described it, he added, “To be 
rediscovered.” Presumably the lack of subsequent collections was a major fac- 
tor when Gray later 1886) dropped his variety macrocarpa, listing B. macrocarpa 
in the synonymy of B. lanuginosa and stating that Nuttall’s plant “must be this 
or the preceding” [B. tenax (L.) Willd. Seemingly, Gray came to regard Nuttall’s 
plant as likely to have been merely an aberrant form of a previously described 


i 
~ 


species, and eliminated B. reclinata from consideration due to the latter's small 
fruit and typically glabrate leaves. 

A second collection of Sideroxylon macrocarpum would not be made until 
the early years of the 20th Century, when the region of Georgia to which 
Nuttall’s plant is endemic was explored botanically by Roland M. Harper for 
his doctoral dissertation, published as A Phytogeographical Sketch of the 
Altamaha Grit Region of the Coastal Plain of Georgia (Harper 1906).“Altamaha 
Grit” refers to a stratigraphic unit now called the Altamaha Formation (Hud- 
dleston 1988). Harper cited Harper 1851 under Bumelia reclinata, and listed 
additional sites from sandhills or dry pine barrens in Tattnall, Montgomery, 
and (present day) Wheeler Counties. The latter were presumably sight records, 
as no specimens were cited. Given their geographic location and habitat, 
almost certainly also represent S. 


” 


Harper's three sight records of “B. reclinata 
macrocarpum. Another collection is known from the first half of the 20th cen- 
tury, Mrs. J.N. Henry 1019 (NY), from 1937. Its label bears no determination ex- 
cept for the word Bumelia, handwritten anonymously just above the label, and 
was otherwise never annotated. 

The next three collections of Sideroxylon macrocarpum known to me are 
at GA: Duncan & Hardin 14622 (in 1952, Toombs County), Broughton s.n. 1957, 
Tattnall Co.), and Cypert 211 959, Pierce Co.). Wilbur Duncan identified his 
and Broughton’s specimens as Bumelia reclinata, Cyperts as B. lanuginosa. These 
specimens were annotated by Cronquist in 1970, two of them simply as “Bumelia 
thornei Crong.” Cronquist elaborated slightly in his annotation of Duncan & 
Hardin 14622: “An unusual specimen which may perhaps best be associated 
with B. thornei.” The habitat information supplied on the labels indicated that 
all three collections came from dry, sandy habitats. That they came from such 
habitats presumably posed no problem for Cronquist, since in naming B. thornei 
he had (erroneously) cited the habitat of the type (Thorne 7345, GA) as “dry 
[my emphasis, here and in the next quotation] live oak woods by cypress swamp” 
(Cronquist 1949). However, this conflicts with the habitat given by Robert 
Thorne himself (1949) for the type locality, namely “moist live oak woods at 


— 


ALLISON 253 


edge of cypress swamp.” Furthermore, Thorne (1949, 1954) gave the habitat of 
the species as a whole as “sandy, moist, open meadows or woods.” Cronquist’s 
mis-annotations of 1970 would have significant consequences 

The paucity of occurrences known for Bumelia thornei was undoubtedly 
the primary reason for its listing, in August 1977, as an endangered species un- 
der provisions of the Georgia Wildflower Preservation Act (Patrick et al. 1995). 
A month later McCollum and Ettman (1977) published a treatment of all the 
species then protected under that statute, witha description, Georgia range map, 
and line drawing of each. This government publication was made available free 
of charge and had wide distribution within the state. In it B. thornei was char- 
acterized as “a small, thorny shrub up to 1.5 m tall,” that was found “in dry, live 
oak woods and scrub oak sandhills.” Clearly their concept of B. thornei derived 
from Cronquist’s original, somewhat faulty description and from at least one of 
his 1970 annotations, of Broughton s.n., whose label gave the habitat as “scrub 
oak sandhills.” 

For more than ten years following McCollum and Ettman 1977, Bumelia 
thornei was considered by most botanists in the Southeast to be a low shrub of 
dry habitats. For example, in 1985 Robert B. McCartney reported to the Georgia 
Department of Natural Resources (GaDNR) his finding of a “new Bumelia 
thornei location,” in Tattnall County (correspondence in files of the Natural 
Heritage Program of the GaDNR). In March of 1987 McCartney leda small group 
of botanists, myself included, to this population, which corresponded perfectly 
in habit and habitat to the treatment of B. thornei in McCollum and Ettman 
1977. At this point Nuttall’s B. macrocarpa had been languishing in synonymy 
for just over a century and completely forgotten by Georgia botanists. 

Not surprisingly, knowledge gained in the decades subsequent to Clark 1942 
and Cronquist 1945 supports a taxonomy of this group of species somewhere 
between the extremes represented by those two workers. To be fair to Cronquist, 
he was frank that the firmness of his taxonomic judgments was proportionate 
to the number of herbarium specimens available to him. The taxa named post- 
Michaux were usually represented by few collections. For some of these, rather 
more material has of course proved helpful, especially when combined with 
significant field experience with the plants, which neither Clark nor Cronquist 
could claim. 

In contrast, Robert Godfrey had considerable experience with the group as 
living plants. For example, he is credited with the earliest known collection of 
Sideroxylon thornei from Florida, made in 1982 (Anderson 1988). Informed by 
his field experience with two taxa that had been treated by Cronquist (1945) as 
Bumelia reclindata var. rufotomentosa (Small) Cronq. and B. tenax f. anomala 
(Sarg.) Cronq., Godfrey (1988) restored them to species rank, as B. rufotomentosa 
Small and B. anomala (Sarg.) Clark. In this work Godfrey added new precision 
to their descriptions and discussed them in some detail, with line drawings of 


254 BRIT.ORG/SIDA 22(1) 


each by Melanie Darst. Both of these buckthorns, however, had specific epithets 
that were preoccupied in Sideroxylon, so their eventual transfer to that genus at 
the rank of species required new names: S. rufohirtum Herring and Judd (1995) 
for B. rufotomentosa, and S. alachuense L.C. Anders. (Anderson 1997) for B. 
danomala. 

In the same work, Godfrey revised the description of Bumelia thornei, giv- 
ing the normal height (accurately) as around 2 1/2 meters, with a maximum 
plant height of 6 meters. He also described its fruits as larger than those of B. 
lanuginosa, whereas Cronquist (1949) in the B. thornei protologue had errone- 
ously stated precisely the opposite.’ Godfrey also correctly made B. thornei a 
wetland plant, giving the habitat as “woods bordering ponds (and creeks?), ap- 
parently where some surface water stands during wet seasons.” 

In 1988 McCartney observed Bumelia thornei at its type locality in Early 
County, Georgia (note in files of Georgia Natural Heritage Program) and, as his 
long, handwritten annotation on a 1985 collection (Rayner & McCartney 2401, 
USCH) makes clear, he realized that the often tall shrub of wetland habitats 
upon which B. thornei was founded could hardly be synonymous with the low 
plant of well-drained, sandy habitats portrayed in McCollum and Ettman 1977. 

Angus Gholson showed me to the type locality of Bumelia thornei in May 
of 1990. Later that year Ifound a number of additional localities for genuine B. 
thornei, including several county records (Anderson 1996). Unaware of 
McCartney’s observations, | came to the same conclusion, that B. thornei was 
not conspecilic with the low plant of dry places in southeastern Georgia. The 
following year | provided a corrected description of B. thornei, in an update of 
Georgia’s Protected Plants McCollum & Ettman 1991), as a tall shrub of wet- 
land habitats, known in Georgia (then) only from the southwestern part of 
state. 


the 


This begged the question of what to call the low, large-fruited plant of dry, 
sandy habitats of southeastern Georgia. The com a of large fruit and com- 
paratively small leaves with patchy, persistent hairs beneath, as well asthe habit 
and habitat, invited comparison with one of the taxa restored to species status 
in Godfrey 1988: Bumelia rufotomentosa [Sideroxylon rufohirtum], considered 
to be endemic to north peninsular Florida. For example, McCartney's 1988 type- 
written annotation of D.A. Rayner & R. McCartney 2401 (USCH) states, “The 
Tattnall County material most resembles B. rufotomentosad from central Florida 


The treatments of Small (1900), Clark (1942), Cronquist (1945, 1949) and others based almost exclusively on 
herbarium collections contain errors regarding (mature) fruit size; sometimes the aie limits are given as too 
large, apparently based on the inclusion of misidentified material, sometimes too small, due to misjudging the 
state of maturity of unripe fruit on older, discolored specimens. Small, for example, described his Bumelia 
rufotomentosa (Sideroxylon rufohirtum) as having slightly smaller fruits than B. reclinata, when in fact they are 


distinctly larger at maturity 


ALLISON 255 


which occurs in similar habitat but has more orbicular, persistent leaves. May 
represent a new taxon!” 

In the early 1990s Carl Nordman, during surveys of rare species and natu- 
ral communities of the Altamaha River basin for The Nature Conservancy of 
Georgia, found several new localities for the seemingly anonymous Sideroxlon 
in Appling, Pierce, Tattnall and Toombs Counties; vouchers for eight of these 
have been cited herein. | also maintained an interest in the plant, looking for it 
whenever my work for the GaDNR took me to the southeastern part of the state 
and making the first collections from Candler and Treutlen Counties. In 1996- 
97, | conducted for the GaDNR, with funding from the US. Fish and Wildlife 
Service, a status survey of Sideroxylon thornei and of the low plant of sandy 
places that had been mistakenly included within it (Allison 2000). Soon there- 
after | shared my findings and some specimens with an authority on 
Sideroxylon, Loran Anderson. Informally, in an abstract, we indicated our in- 
tention to name the plant S. duncanorum (Allison & Anderson 1998). The pro- 
posed specific epithet was to commemorate a lifetime of service to southeast- 
ern botany on the part of the late Wilbur H. Duncan and of his wife and work 
partner, Marion B. Duncan. I plan, instead, to name another Georgia endemic 
shrub (of the Lamiaceae) in their honor. 

While working up the manuscript for the “new species” of Sideroxylon, | 
came across the name Bumelia macrocarpa Nutt. among synonyms of B. 
reclinata listed in Clark 1942. Nuttall’s choice of an epithet meaning “large- 
fruited” demanded further inquiry, as this name would be very appropriate for 
the Georgia endemic. Furthermore, | recalled that Nuttall collected the type of 
Arenaria brevifolia Nutt. ex Torr. @& A. Gray [=Minuartia uniflora Walt.) Matt] 
from Tattnall County, Georgia (Harper 1904), placing him in the vicinity of 
extant locations of the supposedly undescribed Georgia buckthorn. 

In May of 2005 | examined the only sheet (Fig. 1) of Nuttall’s Bumelia 
macrocarpa at the Academy of Natural Sciences, Philadelphia (PH). It bears five 
fragments of Sideroxylon and three paper labels: (1) a primary label that can be 
no older than 1878 (19 years after Nuttall’s death), as it identifies the sheet using 
Gray’s combination, B. lanuginosa var. macrocarpa; (2) Clark’s 1940 annotation 

label, identifying the sheet as B. reclinata; and (3) pierced by one of the frag- 
ments, a scrap with two handwritings. On the latter was written in pencil: “B. 
[blank space for a specific epithet]. Fruit large as a small date! Eatable. Geo.” Ac- 
cording to James Lendemer of PH (pers. comm.), Charles Pickering, a curator of 
PH and close friend of Nuttall’s (Graustein 1967) probably assembled and 
mounted this material from Nuttall’s disorganized and voluminous material at 
PH, and supplied the annotations in black ink on Nuttall’s scrap of a label: the 
specific epithet “macrocarpa Nutt., the quotation marks surrounding the origi- 
nal penciled text, and the identification of the penciled handwriting as Nuttall’s. 
This label is unquestionably Nuttall’s, as it conforms exactly to his practice in 


ee 


256 BRIT.ORG/SIDA 22(1) 


this regard, as described by Graustein (1967). Only the fragment piercing this 
label, as discussed near the end of the introduction, is a match for Nuttall’s de- 
scription of B. macrocarpa and tor the specimens cited above, under additional 
collections examined. Although Nuttall did not specify a type collection, this 
specimen is housed in the institution where he worked for many years, both 
before and after the presumptive collection date of 1830. Therefore I designate 
the fragment at upper left as the lectotype. 

HABITAT 
The habitat of Sideroxylon macrocarpum is sandy, well-drained, and dry-mesic, 
but apparently not quite dry enough for the partly sympatric Chrysoma 
pauciflosculosa (Michx.) Greene or Ceratiola ericoides Michx. to occur. The great 
majority of occurrences have a well-developed overstory of Pinus palustris P. 
Mill. with fire infrequent enough (or excluded long enough) to permit arbores- 
cent oaks to attain flowering size. The latter may include Quercus margaretta 
Ashe ex Small, Q. incana Bartr, Q. laevis Walt., 9. marilandica Muenchh. and/ 
or Q. hemisphaerica Bartr. ex Willd. 

Sites that seemed otherwise suitable but had been subjected to significant 
soil disturbance (e.g. chopping and bedding) were usually devoid of the plant, 
though occasionally a few survivors could be found. The plant’s growth form, 
with the majority of its biomass beneath the litter layer, seems a clear adapta- 
tion to episodic fires as well as for conserving water. Some perennials are pro- 
moted by root fragmentation due to chopping and other soil disturbances as- 
sociated with currently prevalent forestry practices. Sideroxylon macrocarpum 
clearly does not tolerate these practices well. 

Aside from Pinus palustris and the oaks listed above, the tree most fre- 
quently recorded in association with Sideroxylon macrocarpum was Prunus 
serotina Ehrh. Shrubs or woody vines recorded as common associates of S. 
macrocarpum were Licania michauxii Prance, Morella cerifera (L.) Small, Opun- 
tia humifusa (Raf.) Raf., Gelsemium sempervirens (L.) St. Hil, Toxicodendron 
pubescens P. Mill. and Vitis rotundifolia Michx. Frequent herbaceous associates 
included Arist ida stricta Michx., Baptisia perfoliata (L.) R. Br. ex Ait. [., B. 
lanceolata (Walt.) ElL, Eriogonum tomentosum Michx., Cnidoscolus stimulosus 
(Michx.) Engelm. & A. Gray, Pteridium aquilinum (L.) Kuhn, Tephrosia 
virginiana (L.) Pers.,and Vernonia angustifolia Michx. (data from Allison 2000). 

Exceptions to the preceding habitat characterization include powerline 
rights-of-way and road shoulders, where the plant grows in more exposed situ- 
ations. Here the adverse, drying effects of more sunlight reaching the substrate 
are apparently counterbalanced by a regular mowing or bush-hogging regime. 
The latter must surely have much more impact on taller competitors than on 
the low-growing Sideroxylon, promoting its persistence and increase asa result 
of decreased competition. 


— 


ALLISON 257 


COMPARISONS WITH RELATED TAXA 


Whether called Bumelia or Sideroxylon, the temperate species of the southeast- 
ern states have long been considered a natural group. Pennington (1990) states 
that “they are easily distinguished from all Neotropical species by the combi- 
nation of fascicled leaves with conspicuously finely reticulate tertiary vena- 
tion, short-styled flowers, small fruit, and the seed with a bipartite scar.” Re- 
flecting this, the five such species recognized by him (S. lanuginosum, S. 
reclinatum, S. thornei, S. tenax, S. lycioides) are separated, under the first cou- 
plet, from all others in his key to the species of Sideroxylon. More than century 
earlier, Gray (1878, 1886), treating all of these taxa except the then-undescribed 
thornei, emphasized the deciduous leaves and “staminodia nearly as large as 
the proper [medial segments of the] corolla-lobes” as unifying characters. 

At one time or another, plants of Sideroxylon macrocarpum have been con- 
fused with each of the abovementioned species except S. tenax and S. lycioides. 
In fact, as discussed earlier, it is most similar not to any of these but to a less 
well known taxon completely ignored by Pennington, the one now knownas S. 
rufohirtum. 

Sideroxylon tenax and the closely related S. alachuense stand apart from 
all these species most conspicuously by their densely sericeous-tomentose lower 
leaf-surfaces, the pubescence mostly tawny or rufous in the former, mostly sil- 
very in the latter. Sideroxylon lycioides differs from all except S. reclinata s. str. 
in having leaves that are normally glabrate beneath; the former has consistently 
larger leaves and fruit than the latter. Sideroxylon macrocarpum is readily dis- 
tinguished from these two taxa by the leaves persistently, if rather sparsely, stri- 
gose beneath and by occurring in better drained, sandier habitats than is nor- 
mal for them. However, populations of S. reclinatum from south peninsular 
Florida, described (Whetstone 1985) as Bumelia reclinata var. austrofloridensis 
Whetstone, are more similar to S. macrocarpum in having persistent leaf-pu- 
bescence; but the small fruit size (< 9 mm long) and staminodes equaling or 
exceeding the corolla-lobe appendages, indeed indicate closer affinity to S. 
reclinatum s. str. than to S. macrocarpum. 

It seems unlikely that Gray had ever seen material of Bumelia macrocarpa 
when he reduced it to a variety of B. lanuginosa, but he probably felt that the 
persistent foliar pubescence emphasized by Nuttall ruled out synonymizing it 
under B. lycioides or B. reclinata. Perhaps Nuttall’s use of the term “lanuginous” 
to describe the vestiture influenced Gray to place it with B. lanuginosa and not 
B. tenax. Sideroxylon lanuginosum is clearly a heterogeneous taxon, especially 
as regards coloration, density, and persistence of foliar pubescence, with three 
subspecies accepted by Pennington (1990). Only S. lanuginosum s. str. is known 
from Georgia or Florida, however, and like S. tenax, S. lanuginosum subsp. 
lanuginosum has lower leaf surfaces witha persistent pubescence dense enough 


258 BRIT.ORG/SIDA 22(1) 


to obscure all or most of the laminal surface, while in S. macrocarpum and S 
rufohirtum the hairs are much sparser, the majority of the surface always vis- 
ible once the leaves are fully expanded. Among the other subspecies of S 
lanuginosum, only subsp. rigidum (A. Gray) Pennington invites comparison 
with S. macrocarpum, by virtue of its small leaves witha foliar pubescence some- 
times resembling that of the Georgia endemic. The fruit of S. lanuginosumsubsp. 
rigidum is unknown to me; however, this plant of the southwestern U.S.A. and 
northern Mexico is a “shrub or small tree to 8 m” (Pennington 1990). 

No doubt the similarity of foliar pubescence was largely responsible for 
Cronquist’s confusion of specimens of Sideroxylon macrocarpum with S.thornet. 
While these species do share a similar pubescence and comparatively large fruit 
size, they exhibit major differences in habit and habitat. Sideroxylon thornet is 
a larger-leaved (to as much as 13.9 cm long [Anderson 1996)), erect and poten- 
tially tall shrub or small tree of wetlands, usually associated with Taxodium, 
whereas S. macrocarpum is a small-leaved (rarely to 5.2 cm), sprawling shrub 
less than a meter tall, adapted to well-drained upland woodlands dominated 
by Pinus palustris and various species of Quercus L. 

Asstated previously, Sideroxylon macrocarpum is more similar in habit and 
habitat to the allopatric S. rufohirtum than to any other species. They differ in 
several minor respects and in at least two major ones, androecium (including 
staminode) size and seed coloration, as indicated in Figure 2 and Table I. 
re known ranges of Sideroxylon macrocarpumand S. rufohirtumare sepa- 
rated by roughly 190 km (120 miles). Most of this intervening territory is in- 
hospitable to species requiring well-drained soils, as it consists (or consisted, 
prior to human alteration) largely of lowlands, wetlands and bodies of water, 
most conspicuously the Okefenokee Swamp. A change in broad-scale landscape 
within the gap in the two ranges is apparent at once from an examination of a 
Georgia landcover map (Canalos & Clover 1993), in which the most abundant 
cover class assigned throughout the range of S. macrocarpum is “cultivated /ex- 
posed earth.” South of this region the Okefenokee is apparent from a glance at 
this map, but with closer study an area more than twice its size immediately to 
its west is also well differentiated. This region remains dominated by forest land, 
reflecting the area’s comparative unsuitability for agriculture, presumably due 
to its being insufficiently well-drained (Christopher Canalos, pers. comm.). 

| consider it likely that these two buckthorns are sibling species, now geo- 
graphically isolated in large part by the development of the Okefenokee and 
lowlands to its west. Although the oldest peat deposits from the Okefenokee 
have been radiocarbon dated at a little less than 7000 years B.P, for many years 
prior to the period of peat accumulation the landscape there was dominated 
by freshwater marshes (Cohen et al. 1984). Because the Okefenokee developed 
ina basin formed during one of the last great transgressions of the Atlantic Ocean 
over southeastern Georgia, during the very late Pliocene or earliest Pleistocene 


oa 
pny 


om) 


iu 


ALLISON ; ND 259 


Taste 1. Morphological comparison of Sideroxylon macrocarpum and S. rufohirtum. 


Sideroxylon macrocarpum Sideroxylon rufohirtum 

Stems usually copiously thorny; galls rarely Stems less thorny; often with galls (Godfrey 
observed (e.g. Allison 12220, 

Leaves u ually narrowly oblanceolate or l lly broadly but i | 
spatulate oblanceolate, spatulate 

Twig and leaf trichomes mostly whitish or Twig and leaf trichomes vu (rusty brown), 
translucent, generally less abundant generally more a 

Pedicels short, 1-3(-5.5) mm Pedicels longer (1.8 i oe )m 

Sepals unequal (outer 2 shorter), pubescence — Sepals + equal in length, pubescence moderate 

arse to moderate to dense 

Corolla (3.1—)3.7-4.8 mm; medial lobe- Corolla 3.8-5.0 mm; medial lobe-segments 
segments broadly obovate, lateral segments relatively narrow, lateral segments broader 
relatively narrow than in macrocarpum 

Style 1.0-1.3(-1.5) mm Style (1.5-)1.7-1.9 mm 

Ovary glabrous or medially villosulous Ovary covered with hairs 

Anthers 1.1-1.3(-1.5) mm, filaments 1-2 mm Anthers 0.7-0.9(-1.0) mm, filaments 1.5-2.1 mm 

Staminodes (1.2-)1.3-1.8(-2.0) mm,reaching | Staminodes (1.7—)2.0-2.5 mm, nearly reaching 
(1/3-)1/2 to 2/3 the length of the medial the length of the medial corolla-lobe 
corolla-lobe segments and exceeded g t ling the | | 
by the lateral segments occasionally with sagittate bases 

Seed evenly brown Seed variegated (brown and yellowish) at 

maturity 


(Rich 1984), the area was unsuited to plants of well-drained soils for at least 
several millennia prior to 7000 years B.P. The similarity of the habitats of 
Sideroxylon macrocarpum and S. rufohirtumd t an alternative expla- 
nation: that their comparatively large fruit and low s stature resulted from par- 
allel or convergent evolution, and not direct descent from a common ancestor. 

Whatever the cause, Sideroxylon macrocarpum has been isolated reproduc- 
tively long enough to consistently exhibit at least one morphological extreme 
within the temperate species-group endemic to the southeastern U.S.A. a 
smaller size of staminode relative to corolla lobe. Since it is further differenti- 
ated from the similar but allopatric S. rufohirtum by having evenly brown 
mature seeds rather than variegated ones, I have no hesitation in recognizing 
the Georgia plants at the level of species. 


CONSERVATION STATUS 


The range-wide decline of Pinus palustris-dominated communities due to fire 
suppression, conversion to other forest types or to farmland, pasture and, in- 
creasingly, residential uses, is too well-documented to require discussion here. 
Whether Sideroxylon macrocarpum as a species has suffered the same degree 
of decline over the last century, or any decline at all, cannot be shown by my 


260 BRIT.ORG/SIDA 22(1) 


observations, due toa lack of baseline data. Since populations seem best devel- 
oped where there is a moderate oak understory, it seems likely that fire sup- 
pression, where not too rigorous or prolonged, has benefited mid-successional 
species like S. macrocarpum, at least in some relatively xeric habitats over deep 
sands. Too short a fire interval, with too-frequent destruction of aboveground 
biomass, outweighs the benefit of reduced competition. The converse: too-pro- 
longed exclusion of fire from the habitat leads to excessive shading from clos- 
ing of the canopy and crowding out by non-fire-adapted competitors. 

Immediate threats to Sideroxylon macrocarpum arise chiefly from conver- 
sion of its habitat to short-rotation “industrial” pine plantation. Impacts from 
current practices include soil disturbance (damage to the deep root system), 
herbicide use (to kill competitors of pine, such as Sideroxylon), and fertilizer 
application. The latter is of no benefit to plants, like this species, that have 
evolved adaptations making them tolerant of nutrient-poor soils and giving 
them a competitive advantage there, while fertilization is distinctly beneficial 
to broadly-adapted, weedy competitors, both native and exotic. 

Another cause for concern is the movement away from mowing asa means 
of maintaining rights-of-way and toward a dependence on herbicides. As dis- 
cussed above, mowing can provide a competitive advantage to low-growing 
species such as Sideroxylon macrocarpum. Herbicide use, by contrast, leads to 
their replacement over time by a few herbicide-resistant species, which would 
seem eventually to require either a return to mowing or changes to the herbi- 
cide regime. 

At present, as the many collections cited above from the last two decades 
attest, the plant is not extremely difficult to find within suitable habitat in the 
part of Georgia to which it appears to be endemic. However, I suggest consider- 
ation be given to amending the list of plants receiving protection under the 
Georgia Wildflower Preservation Act, to replace Sideroxylon thornei with S. 
macrocarpum, for several reasons. For one, as indicated earlier, the apparent in- 
tent in listing B. thornei was to provide protection to the plant now known as S. 
macrocarpum, the result of mistaken specimen annotations by Cronquist in 
1970. Secondly, genuine S. thornei is now known to be much more widespread 
and abundant than was once believed (Patrick et al. 1995; Anderson 1996, Alli- 
son 2000), with a fair number of occurrences on public lands. Furthermore, 
the wetland habitats of S. thornei receive protection under the federal Clean 
Water Act (CWA), without regard to any rare species that may be present. This 
law affords a much greater level of protection to wetland plants than is pro- 
vided by state or federal protected species laws, which provide little or no habi- 
tat protection to plants. The CWA provides no benefit to obligate upland spe- 
cies such as S. macrocarpum, part of the suite of longleaf pine associates so 
famously in decline; it serves instead to channel development away from wet- 
lands and into habitats like those of S. macrocarpum. Finally, I believe that each 


ALLISON, _AFORGOTTEN GEORGIA ENDEM 261 


state wildlife agency should recognize a special responsibility to provide for 
the welfare of those species found nowhere except within the borders of its state. 

Of course, the intention of laws like Georgia’s Wildflower Protection Act is 
to prevent additional species from suffering the same fate as the dodo (Raphus 
cucullatus): extinction. This famously extirpated flightless bird of Mauritius is 
an especially appropriate example because of the link posited by Temple (1977) 
between its eradication and the decline there of a congener of Sideroxylon 
macrocarpum, S. majus (Gaertn. f.) Baehni (syn.: Calvaria major Gaertn. f.). In 
brief, Temple averred that the Sideroxylon was represented by a few trees esti- 
mated to be more than 300 years old, despite the regular production of well- 
formed fruit. He theorized that the supposed absence of reproduction in the 
Sideroxylon was due to the extinction of the dodo, which presumably scarified 
the seeds in its gizzard after ingestion of the fruit and that this was required for 
their germination. He tested this by force-feeding seeds of the Sideroxylon to 
domesticated turkeys, after which he extracted some of the seeds from the scat 
and successfully germinated a few of them.* 

Whether Temple’s hypothesis is correct or not (for a contrary view see 
Owadally & Temple 1979), it seems possible that the comparatively large fruit 
size of Sideroxylon macrocarpum and of its sibling species, S. rufohirtum, re- 
flects natural selection to make the fruit more attractive to another terrestrial 
animal of conservation concern, the gopher tortoise (Gopherus polyphemus). 
The range of S. macrocarpum is nested entirely within the range (mapped in 
Auffenberg & Franz 1982) of this turtle, which is often syntopic with the 
Sideroxylon today (pers. obs.; Fig. 5). It is probable that, prior to the serious de- 
cline of the tortoise, it was a regular associate of the buckthorn in the open, 
well-drained, sandy habitat required by the plant and favored by the reptile. 
The low stature of this shrub certainly places its fruit within easy reach of the 
tortoise, though admittedly this may be a secondary result of selection due to 
recurrent fires that are thought to be required to maintain ideal habitat for both 
species. It must also be admitted that fruit size in Sideroxylon is probably cor- 
related with seed size, and that the greater storage capacity of larger seeds has 
survival value in drought-prone habitats. Frugivory by Gopherus may not have 
been the primary selective force promoting lower stature and larger fruit. But it 
should also be noted that, compared to the species with which they have been 
confused, the large fruits of S.macrocarpum and S. rufohirtum more closely re- 
semble a fruit reported to be commonly consumed by gopher tortoises: that of 
Serenoa repens (Bartr.) Small (Breininger et al. 1986). As in S. macrocarpum and 
its relatives, the fruit of this native palm is a black, fleshy, mostly subglobose 
fruit, most of whose volume is occupied by a single large seed (as figured in Uhl 


4in 1978 this purported case of coevolution of bird and tree was a subject of one of the celebrated series of 
essays by the late Stephen A. Gould, This View of Life,in the popular scientific magazine Natural History. 


262 BRIT.ORG/SIDA 22(1) 


& Dransfield 1987). Birkhead et al. (2005), analyzing gopher tortoise scat, un- 
fortunately from outside the geographic range of either S. macrocarpum or S. 
rufohirtum, reported ingestion by gopher tortoises of the large-seeded fruits of 
several dicotyledonous genera, including Asimina Adans., Licania Aubl. and 
Prunus L.,and found that such seeds were defecated intact. They concluded that 
these animals play an important role as dispersal agents for plants with large- 
seeded fruits in pine savannas of the Southeast. It should also be noted that the 
gap separating the ranges of S. macrocarpum and S. rufohirtum (see discussion 
above, following the table) corresponds roughly toa gap depicted on the gener- 
alized range map of Gopherus polyphemus appearing in Ernst et al. 1994. 

Let us hope that future generations will be able to see—if not the hapless 
dodo—thriving populations of gopher tortoises, big-fruited buckthorns, and as 
wide a spectrum as possible of the rest of the incalculable diversity of life-form 
bequeathed and entrusted to us all. We have the means; it remains to be seen 
whether we also have the will. 


ACKNOWLEDGMENTS 


Partial funding of this project was provided by the United States Fish and Wild- 
ife Service and the Georgia Department of Natural Resources. Particular thanks 
goto Loran Anderson, Professor Emeritus at Florida State University, who helped 
in many ways, including (but not limited to) assistance in locating populations 
of Sideroxylon rufohirtum, loaning mature seed of it from FSU specimens, and 
encouragement to complete the work. lam also grateful to the curators and other 
staff and researchers at other herbaria consulted during this study: Kent Per- 
kins at FLAS; the late Wilbur Duncan and Michael Moore at GA; Emily Wood 
at GH; James Solomon at MO; Jacquelyn Kallunki and Stella Sylva at NY; at PH: 
James Macklin (who also provided oe a of the type specimen), Nancy 
Kahn and, especially, James Lendemer, whose comments prompted a thorough 
examination of the Nuttall material; Daher Bell and Marjorie Knowles at US; 
and John Nelson at USCH. Thanks also to Keith Tassin and Eric Van De Genachte 
of The Nature Conservancy, the former for assistance in visiting populations in 
the Altamaha River basin, the latter for providing me his early-season speci- 
men (ina similar stage as Nuttall’s type; deposited at PH to facilitate compari- 
son). Thanks go to Franklin Snow of South Georgia College for showing me the 
population he found along the Oconee River; thanks to Mr. Snow also for pro- 
viding me with his specimen and to Carl Nordman of NatureServe, who en- 
trusted me with his several collections of the plant. Thanks as well to Greg 
Krakow of the Georgia Natural Heritage Program, who helped with prepara- 
tion of the map. Thanks also for valuable discussions with Robert McCartney, 
Thomas S. Patrick, and Christopher Canalos, and to R. David Whetstone for 
helpful suggestions to improve the manuscript. Finally, I thank José Tallet for 


—_— 


ALLISON 263 


improvements to my Spanish abstract, and Mark A. Garland for correcting my 
(ultimately unneeded) Latin diagnosis for “Sideroxylon duncanorum.” 


REFERENCES 


Auuson, J.R. 2000. Summary report: Section 6 status survey of Sideroxylon thornei, swamp 
buckthorn, in Georgia. Georgia Dept. of Natural Resources, Social Circle. 

Atuson, J.R. and L.C. ANDERSON. 1998. Sideroxylon duncanorum (Sapotaceae), a new buck- 
thorn from Pinus palustris-dominated communities of southeastern Georgia [abstract]. 
ASB Bull. 45:120. 

Anberson, L.C. 1988. Noteworthy plants from North Florida. Ill. Sida 13:93-100. 

Anperson, L.C. 1996. New geographical and morphological data for Sideroxylon thornei 

Sapotaceae). Sida 17:343-348. 

ANberSON, L.C. 1997. Sideroxylon alachuense,a new name for Bumelia anomala (Sapotaceae). 
Sida 17:565-567. 

AUFFENBERG, W. and R. FRANZ. 1982. The status and distribution of the gopher tortoise 
(Gopherus polyphemus). In: R.B. Bury, ed. North American tortoises: conservation and 
ecology. Wildlife Research Report 12, U.S. Fish and Wildlife Service, Washington, DC. 

BiRKHEAD, R.D., C. Guyer, S.H. Hermann, and W.K. MicHener. 2005. Patterns of folivory and seed 
ingestion by gopher tortoises (Gopherus polyphemus) in a southeastern pine savanna. 
Amer. Midl. Naturalist 154:143-151. 

BREININGER, D.R., PA. ScHMaLzer, D.A. Rybene, and C.R. Hinkle. 1986. Burrow and habitat relation- 
ships of the gopher tortoise in coastal scrub and slash pine flatwoods on Merritt Is- 
land, Florida. Florida Game and Fresh Water Fish Commission, Tallahassee. 

Canatos, C.G. and LJ. Clover. 1993. Landcover map of Georgia. J. Urban Regional Inform. 
Systems (URISA) 5:99-103. 

Ciark, R.B. 1942. A revision of the genus Bumelia in the United States. Ann. Missouri Bot. 
Gard. 29:155-182. 

CoHeN, A.D., MJ. ANDREJKO, W. Spackman, and D.Corvinus. 1984.Peat deposits of the Okefenokee 
Swamp. |n:A.D.Cohen, D.J.Casagrande, M.J.Andrejko, and G.R. Best, eds. The Okefenokee 
Swamp: its natural history, geology, and geochemistry. Wetlands Surveys, Los Alamos, 


— 


eo 


Cronauist, A. 1945. Studies in the Sapotaceae, Ill. Dipholis and Bumelia. J. Arnold Arbor. 
26:435-471. 

Cronauist, A. 1946. Studies in the Sapotaceae, Il Survey of the North American genera. 
Lloydia 9:241-292. 

Cronauist, A. 1949. Noteworthy plants of Georgia. Castanea 14:102-108. 

pe CANnDoLte, A. P. 1844. Prodromus systematis naturalis regni vegetabilis. Vol. 8. Treuttel & 
Wurtz, Paris, Strasbourg & London. 

Ernst, C.H., R.W. Bargour, and J.E. Lovicn. 1994. Turtles of the United States and Canada. 
Smithsonian Institution, Washington. 

Goorrey, R.K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Geor- 
gia and Alabama. Univ. of Georgia Press, Athens. 


264 BRIT.ORG/SIDA 22(1) 


Graustein, J.E. 1967. Thomas Nuttall, naturalist; explorations in America, 1808-1841. Har- 
vard Univ. Press, Cambridge. 

Gray, A. 1878. Synoptical flora of North America [1st ed], lvison, Blakeman, Taylor & Co., 
New York. 

Gray, A. 1886. Synoptical flora of North America, 2nd ed., Smithsonian Institution, Wash- 
ington. 

Harper, R.M. 1904. The type-locality of Arenaria brevifolia. Torreya 4:138-141. 

Harper, R.M. 1906.A phytogeographical sketch of the Altamaha Grit Region of the Coastal 
Plain of Georgia. Ann. New York Acad. Sci. 17(1):1-414. 

HerrING, B.J.and W.S. Juop.1995.A floristic study of Ichetucknee Springs State Park, Suwannee 
and Columbia Counties, Florida. Castanea 60:318-369. 

HUDDLESTUN, P.F.1988.A revision of the lithostratigraphic units of the Coastal Plain of Geor- 
gia. The Miocene through Holocene. Georgia Geol. Surv. Bull. 104. 

McCottum, J.L.and D.R. Erman. 1977. Georgia's protected plants. Georgia Dept. of Natural 
Resources, Atlanta. 

McCottum, J.L.and D.R. Erman. 1991 [revision].Georgia’s protected plants.Georgia Dept. of 
Natural Resources, Social Circle. 

Nurtatt, T. 1849. The North American sylva. Vol. 3. Smith & Wistar, Philadelphia. 

Owabalty, A.W. and S.A. Tempte. 1979. The dodo and the tambalacoque tree. Science 
203:1 363-1364. 

Patrick, 1.S.,J.R. Alison, and G.A. Krakow. 1995. Protected plants of Georgia. Georgia Dept. of 
Natural Resources, Social Circle. 

PENNINGTON, T.D. 1990. Sapotaceae. Fl. Neotropica Monogr. 52:1-770. 

PennincTon, T.D. 1991. The genera of Sapotaceae. Royal Botanic Gardens, Kew. 

Rich, F.J. 1984. An ancient flora of the eastern Okefenokee Swamp as determined by pa- 
lynology.In:A.D.Cohen, D.J.Casagrande, MJ.Andrejko, and G.R. Best, eds. The Okefenokee 
Swamp: its natural history, geology, and geochemistry. Wetlands Surveys, Los Alamos, 
NM. 

Smatt,J.K. 1900. The genus Bumelia in North America. Bull. New York Bot. Gard. 1:437-447. 

Smatt, J.K. 1933. Manual of the southeastern flora. Univ. of North Carolina Press, Chapel 
Hill. 

Tempte, S.A. 1977.Plant-animal mutualism:coevolution with dodo leads to near extinction 
of plant. Science 197:885-886. 

THorne, R.F. 1949. The flora of southwestern Georgia. Ph.D. thesis, Cornell University. 

THorne, R.F.1954. The vascular plants of southwestern Georgia. Amer. Midland Naturalist 
52:257-327. 

Uri, N.W. and J. Dransrieto. 1987. Genera palmarum. Allen Press, Lawrence, KS. 

Wuerstone, R.D. 1985. Bumelia reclinata var. austrofloridensis (Sapotaceae), a new variety 
from South Florida, U.S.A. Ann. Missouri Bot. Gard. 72:544-547. 

Woop, C.E., JR. and R.B. CHANNeLL. 1960. The genera of the Ebenales in the southeastern 
United States. J. Arnold Arbor. 41:1-35. 


SEXUALAND APOMICTIC PRAIRIE FEEABANE 
(ERIGERON STRIGOSUS) IN TEXAS: GEOGRAPHIC 
ANALYSIS AND A NEW COMBINATION 
(ERIGERON STRIGOSUS VAR. TRAVERSII, ASTERACEAE) 


Richard D. Noyes, Heidi Gerling, Carla Vandervoort 


Department ane and Evolutionary Biology 
sity of Colorado 
a aa 80309, U.S.A. 


ABSTRACT 


] ] - ft] 1 


he ae oe (sexual diploid vs 
vit Id.) in Texas was estimated through the cles of pollen quality for Rerbaniinis sped: 
mens ae ae oe Research Institute of Texas (BRIT) and the University of Texas (TEX). The 
approximate geographic coordinates ad 285 una cousculens were eels nes a eroprapnic 


distributions of sexual vs. af i I 

software. The specieaa as a whol I the eastern hall a Texas, ainouey one one 
specimen I | Pande AT i were far 

sexual plants (245 vs. 40 specimens), and logi litud or 

ral area, precipitation, and soil. Sexual plants were ced to Piney wee ve Pee Oak Sash 
habitats in extreme eastern Texas, and correspond to Erigeron traversii Shinners. This ta 
described in 1964 but ea included as a synonym of E. strigosus. This ee ne 
is similar to that recorded for other taxa that include sexual and apomictic populations. The new 


combination E. strigosus var. traversii (Shinners) Noyes is proposed. 


RESUMEN 
El modo cc peo ea pe VS. ad ane de Erigeron strigosus Muhl. ex 
Willd.e especimenes de herbario del 
es Research Institute de Texas (BRIT) y de la Unnersead de Texas oe Se determinaron las 
J i] at : ees Geers 1 : ate 


oO ft Oo Oo 
de pl l icti un eee de Sistemas de Informacion G 
(GIS). La especie en conjunto esta Pon la me este de oes apace se registré un schecinen adel 
norte de Panhandle. I las sexuales (245 vs. 


p 
40 ee y mostraron mayor amplitud Heme con aan al area p natural precipitacion, 
] t 


y suelo restringidas a | 


este gs: Texas, y So a aoe traversii Suna Este taxon se EGeSOUDIO en es pero 
E. st 


rigosus. El 


L fo) (=) 5 


c 
] t t incl blaci 1 Feecmicn ee S I | mbinacion E 


L x 
strigosus Var. traversii (Shinners) Noyes. 


Many plant taxa in temperate regions include both sexually and apomictically 
reproducing populations (Gustafsson 1946-1947; Grant 1982; Asker @ Jerling 
1992). These taxa occur predominantly in the Asteraceae, Rosaceae, and Poaceae, 
and include diplosporous apomicts (the unreduced gametophyte arising di- 


SIDA 22(1): 265 — 276. 2006 


266 BRIT.ORG/SIDA 22(1) 


rectly from divisions of the megaspore mother cell) as well as aposporous 
apomicts (the unreduced gametophyte arising from a vegetative cell of the 
ovule). Apomictic taxa are almost always polyploid, and though they produce 
genetic clones through maternal function, they also usually produce reduced 
pollen that can function in crosses with facultative apomicts and sexual plants. 
The latter are frequently restricted to the diploid level. Apomictic populations 
and sexual populations may have different distributions, a pattern that is re- 
ferred to as geographic parthenogenesis. Apomicts, relative to sexuals, have 
larger ranges, range into higher latitudes, range to higher altitudes, and have a 
greater tendency to colonize once-glaciated areas (Bierzychudek 1987). 

In plants, pollen quality is highly diagnostic for mode of reproduction 
(Gustafsson 1946-1947). Sexual plants typically produce pollen of uniform size 
and high viability as estimated with pollen vital stains. Apomictic plants, on 
the other hand, produce pollen that is of relatively low quality, with high varia- 
tion in size, a high proportion of aborted grains, and often including miniature 
aborted pollen grains (i.e., micrograins). These distinguishing features are readily 
observed from pollen samples obtained from fresh pollen samples as well as 
from herbarium specimens. 

Erigeron strigosus Muhl. ex Willd., the prairie fleabane, is abundant in the 
United States from the Atlantic coast to the Great Plains and occurs sporadi- 
cally in the West (USDA, NRCS 2005). The species has been considered to be 
uniformly apomictic in monographs (Cronquist 1947) and floristic treatments 
(e.g., Barkley 1986; Hickman 1993). Diplospory was documented cytologically 
(Holmgren 1919) and most of the published chromosome number counts for 
the species are triploid (2n = 27; 14 / 38 counts = 37%), tetraploid (2n = 36; 6 / 38 
counts = 16%), or hexaploid (2n = 54; 4 / 38 counts = 11%) (Compiled from Index 
to Plant Chromosome Numbers, 1955-2000). Nesom (1978), however, based on 
diploid counts (2n = 18) from Alabama, Louisiana, and Florida, considered the 
strong likelihood of sexual populations of E. strigosus in the southeastern United 
States. Sexual reproduction was subsequently verified cytologically for diverse 
diploid populations in the southeastern United States (Noyes & Allison 2005). 
Sexual diploid populations included E. strigosus var. calcicolaJ. Allison, princi- 
pally restricted to cedar glades in Tennessee and Alabama, E. strigosus var. 
dolomiticola J. Allison, restricted to a single dolomite glade in Bibb Co., Ala- 
bama, and plants occurring on the piedmont and coastal plain of South Caro- 
lina, Georgia, Florida, and Alabama. In contrast to apomicts, all sexual popula- 
tions investigated were characterized by tetrasporic ovule development. 

The coastal plain is continuous from the Atlantic states to east Texas, and 
although Erigeron strigosus is abundant throughout, biosystematic analysis has 
been wanting. In addition, E. traversii Shinners was described to include sub- 
tly distinct plants in the Pineywoods of Texas and adjacent Louisiana that flow- 
ered earlier than typical E. strigosus (Shinners 1964). Although accepted in the 


NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 267 


Manual of the Vascular Plants of Texas (Correll & Johnston 1970), the taxon 
was later provisionally listed as a synonym of E. strigosus owing to the absence 
of conspicuous morphological characteristics Johnston 1990). That precedent 
has been followed in checklists (e.g., Kartesz 1992; Hatch et al. 2001; USDA, NRCS 
2005) and floras (Flora of North America, Asteraceae 2006, G.L. Nesom, BRIT, 
pers. comm.). 

The objective of this work was to investigate the occurrence of sexual vs. 
apomictic plants of E. strigosus in Texas based on analysis of herbarium speci- 
men pollen samples. Further, detailed geographic distribution and phenologi- 
cal patterns were analyzed, as was the relationship between reproductive mode 
and those plants previously identified as E. traversii. 


METHODS 


Specimens were loaned to the University of Colorado (COLO) from the Botani- 
cal Research Institute of Texas (BRIT) and the University of Texas (TEX). Pollen 
was removed from each specimen to estimate mode of reproduction. Five to ten 
florets that had newly opened or were 1-2 days (estimated) from anthesis were 
removed from single capitula for each specimen. Florets were placed ona stan- 
dard microscope slide with ca. 15 ul Cotton blue in Lactophenol (Stanley & 
Linskens 1974) and allowed to saturate 2 days—1 week. Pollen was released from 
the softened florets as necessary by tearing a longitudinal slit into the corolla 
tubes using dissecting needles. Floral debris was removed, a cover slip was ap- 
plied, and pollen was allowed to stain for ca. 1 week prior to observation. The 
stained pollen was evaluated visually using bright-field microscopy at 400x. 
A pollen sample was identified as being produced by a polyploid apomictic plant 
if it included a high proportion of aborted grains, large grains (usually some 
grains > 17 um diam), and / or micrograins. Conversely, a pollen sample was 
identified as being produced by a sexual plant if it generally included a low 
proportion of aborted grains, was uniformly small (ca. 12-15 um diam), and 
lacked micrograins. Measurements of grains, as required, were performed on 
pollen images captured and analyzed using AnalySIS (v3.1; Soft Imaging Sys- 
tem, GmbH 1989-2001). 

The longitude and latitude of each collection location was determined from 
the data provided on the herbarium specimen label. This was accomplished 
initially using the mapping software TOPO! Texas (Copyright 2004, National 
Geographic Maps, Evergreen, CO) and later using TOPO USA (version 5.0, 
DeLorme, Yarmouth, ME), facilitated on occasion with text web inquiries ae 
Google (Google, Inc, Mountain View, CA). Locality data were compiled in Excel 
(version 10.0.4, Microsoft Corp.). If not explicitly stated, distances and direc- 
tions from towns given on herbarium specimen labels were assumed to lie along 
existing roads. In most cases, the collection data were sufficient to estimate co- 


268 BRIT.ORG/SIDA 22(1) 


ordinates with a high degree of confidence. For a few older specimens, only a 
county or town was provided, in which case coordinates were selected at the 
geographic center of the county or town. To evaluate differences in the phenol- 
ogy of apomictic vs. sexual plants, collection dates were compared statistically 
using Excel. This test assumed that the herbarium specimen collection dates, 
taken as a whole, accurately reflected the duration of flowering in the species. 

Localities were mapped using Geographical Information System (GIS) soft- 
ware ESRI ArcInfo (v. 9.0, copyright 1999-2004; http://wwwesri.com). Dupli- 
cate speci within or between herbaria were removed from the database so 
that subsequent analyses were performed only on unique collections. Asa check 
on locality coding, county names for data plotted in ArcInfo were extracted and 
compared with the original data set. In addition, ca. 20% of the herbarium speci- 
mens were selected at random and plotted localities were evaluated for accu- 
racy. Geographic distributions of sexual and apomictic plants were analyzed 
visually and statistically with respect to rainfall, natural areas, and soils using 
GIS data layers for Texas available from the Texas General Land Office (http:// 
www.glostate.tx.us/gisdata), the Texas Park and Wildlife Department (http:// 
www.tpwd.state.tx.us/landwater/land/maps/gis/map_downloads), and the 
United States Department of Agriculture (http://www.ncgce.nrcs.usda.gov/ 
products/datasets/statsgo/data/tx.html), respectively. The soil classification 
employed followed the taxonomy proposed by the United States Department of 
Agriculture that recognizes 12 soil orders and 64 soil suborders (Soil Survey 
Staff 1999). Specific environmental point data for collections were obtained 
using the Intersect Point Tool (Beyer 2004), and statistical analyses were con- 
ducted using Excel. 


RESULTS 


A total of 311 Texas collections were studied. These included 184 specimens from 
BRIT and 127 specimens from TEX. In addition, floral material from the holo- 
type of Erigeron traversii Shinners (C.L. Lundell 11,093, 11 Apr 1942, BRIT) was 
obtained via special arrangement with curators (185 BRIT Texas collections 
analyzed, 312 collections total). Twenty collections were duplicated between 
BRIT and TEX, one collection was represented by two separate sheets at TEX, 
and three collections were represented by two separate sheets at BRIT. In total, 
there were 288 unique collections between the two institutions. Three collec- 
tions (1%), all from TEX, were excluded from subsequent analyses: G.L. Fisher 
38,018 (1938) and W.R. Carr 11,924 (1992) lacked capitula at a suitable age from 
which pollen could be obtained, and the locality (even to county) could not be 
determined for A. Gieseuschlag s.n. 1928). The earliest collection (FE Lindheimer 
6978) was made near Austin in 1844, and the largest portion of specimens (113, 
40%) was collected in the 1940s (Fig. 1). Although there were specimens pre- 
pared by 95 principal collectors, the majority (52%) was collected by just seven 


NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 


269 
120 
100 a 
“” 
O 
ss 80 
iS) 
2 
© 60 
O 
fe) 
= 40 
ji EL | 
0 +=; T T T T Tt .. T T T LLL, 
— —_ — —_ = —_ oh — —_ — =k ah — hk = = ie) 
o7on7n ow Mm oOo mo 6 6 6 6 © © 6 © © © 6 
khaonntowoeosz NY © FB aAON © © GS 
a — = — 
0 fF DH DH HF HH HF HHH HG H 
Fic. 1. Erigeron strigosus collections by decade in Texas. Solid b ictic (n = 245) b I (n = 40). 


r 
7 
es 
r 
c 
: 
: 
s 
< 
< 


botanists: L. Shinners, 40 (14%); E. Whitehouse, 29 (10%), V.L. Cory, 27 (9.5%), 
DS. Correll, 13 (5%); B.C. Tharp, 13 (5%), and CL. Lundell, 11 (4%), who were 
prolific in the middle decades of the 20th century, and WR. Carr, 14 (5%), whose 
collections were prepared in the 1990s. 

The 285 specimens were collected in 87 of 254 Texas counties (Fig. 2A), 
which are concentrated in north central and east Texas. Most of the collections 
(87%) were restricted to Pineywoods (88, 31%), Post Oak Savannah, (87, 31%) 
and Blackland Prairie (73, 26%) (Fig. 2B). The remaining 37 collections (13%) 
occurred in adjacent regions to the west (Edwards Plateau, 12; Rolling Plains, 
14; Llano Uplift, 4) or to the south (Gulf Coast Prairies and Marshes, 5; South 
Texas Brush Country, 1). Additionally, one specimen occurred in the extreme 
north Panhandle in Lipscomb County. It was the only specimen to occur in the 
High Plains region of the state. No specimens were collected in the Trans Pecos 
region or Coastal Sand Plain of southern Texas. The western boundary of Eri- 
geron strigosus in Texas occurs abruptly at ca. 99°W in the central part of the 
state. This corresponds approximately to the 28 in. precipitation isohyet (Fig. 
2C). The Lipscomb county outlier is exceptional in longitude and rainfall, oc- 
curring at LOO’W in a region with approximately 23 in. annual rainfall. 

Of the 285 specimens, 40 (14%) were determined to possess pollen charac- 
teristic of sexual diploids, and 245 (86%) were determined to possess pollen 
characteristic of polyploid apomicts (Fig. 3). Polyploid apomicts through- 
out the range of the species in Texas. Sexual diploid plants, however, were re- 
stricted to Pineywoods (34 / 40, 85%) and Post Oak Savannah (6 / 40, 15%). 


270 BRIT.ORG/SIDA 22(1) 


oO sexual 


A apomictic 


precipitation. Isohyet values in inches. Scale bars = 100 km. 


Sexual plants were collected in 17 counties within these two regions. In six of 
these counties, only sexual plants have been collected. However, in the other 11 
counties, both sexual and apomictic plants have been collected. The oldest 
sexual specimen was collected by CLL. York in 1928 (s.n., Gregg Co.). There is no 
significant trend in the proportion of sexual plants collected by decade in the 
20th century (r = 0.19, P = 0.58). This indicates that both sexual and apomictic 


NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 271 


Fic. 3. Pollen from herbarium specimens of Erigeron strigosus. A. The holotype of E. traversii (CL. Lundell 11,093, 
Nacogdoches Co., BRIT), sexual. B. L.H. Shinners 32,543 (Morris Co., BRIT), apomictic. Arrow indicates a micrograin. Scale 
bar = 20 pm. 


plants have coexisted in Texas since at least the 1920s, and that their relative 
proportions have not changed dramatically. The holotype of Erigeron traversii 
was determined to have pollen characteristic of sexual diploids, as did the speci- 
mens L. H. Shinners 7618 (BRIT), 18,387 (BRIT), 18,402 (BRIT), 18,450 (BRIT), 
18,503 (BRIT), and B.L. Turner 4377 (TEX), which were cited in the original de- 
scription of EF. traversii (Shinners, 1964). E. Whitehouse 20,861 (BRIT), the eighth 
specimen cited by Shinners for Texas, was not present among the loan materials. 

Precipitation at sites where sexual plants were collected ranged from 41 to 
57 in. / year (mean = 48.2 in., SD = 4.9), while values at apomictic sites ranged 
from 23 to 57 in. / year (mean 39.1 in., SD = 6.3). Precipitation values for apomic- 
tic plants, though overlapping, were significantly lower than those for sexual 
plants (Student t, P = 6.04E-17). Sexual plants predominantly occurred on 
leached forest soils with subsurface clay (alfisols, 18 collections; ultisols, 18 col- 
lections). Four sexual collections occured on poorly developed entisols. Apomic- 
tic plants in Oak Savannah and Pineywoods regions also occurred on alfisols 
(72 collections) and ultisols (28 collections), but also were found on clay-rich 
vertisols (11 collections), and dark grassland mollisols (12 collections). Six 
apomictic collections corresponded to entisols. 

Collections dates of specimens of Erigeron strigosus (Fig. 4) indicate that 
flowering commences in March, peaks in April and May, and continues spo- 
radically at reduced levels until October or early November. One exceptional 
apomictic specimen (R,J. Fleet wood 10,050, Nacogdoches Co., TEX) was collected 
5 Jan 1972 (the collector noted the presence of ice on the plant). The average 
collection date for sexual plants (for 39 collections made from March to June) 
was 22 April (SD = 23.3 days). The average collection date for 210 apomictic 
plants made during the same time period was significantly later (10 May, SD = 


272 BRIT.ORG/SIDA 22(1) 


| q | | 
q . . | 
‘| apomictic 
= 
} 


C] sexual 


Relative Proportion 


T 
| 
1 
T 
| 
t 
I 
T 
I 
qT 
I 
rn 
T 


t 
| 
A 
+ 
| 
t 
| 
qT 
| 
t 
| 
1 
T 
I 


JtrF>m{ta};{m{si fila 


Fig. 4. Erig eu? W * oo | JIi+,f £ } } + + 7b 


22.5 days, Student t, P = 2.5E-06). The same tendency was observed for asexual 
plants occurring in only those counties where sexual plants also occurred (n = 
22, avg. = 9 May, SD = 20.4 days; Student t, P = 0.006). 


DISCUSSION 


Sexual and apomictic Erigeron strigosus in Texas 
The greater range of habitat, precipitation, and tolerated by apomictic Eri- 
geron strigosus in Texas is consistent with the hypothesis (Baker 1965; Lynch 
1984) that asexual lineages, through polyploidy, hybridization, and genome level 
selection, evolve general-purpose genotypes that are capable of exploiting di- 
verse habitats. However, this pattern is al nsistent with the hypothesis that 
apomictic reproduction may be favored in fragmented marginal habitats be- 
cause it circumvents the deleterious effects of inbreeding depression and mate 
scarcity (Haag & Ebert 2004). Detailed, fine-scale analysis of populations, es- 
pecially in regard to population size, habitat, and disturbance, will be required 
to differentiate between these two hypotheses. Superficially, the geographic 
pattern for E. strigosus is similar to that for Taraxacum officinale in Europe. 
Sexual diploid populations of that taxon are restricted to low-elevation habi- 
tats in south central and southeastern Europe, while apomictic populations are 
found throughout northern Europe (Den Nijs @ Menken 1994). 

The historical record indicates that sexual and apomictic Erigeron strigosus 
have co-existed in Texas at least since the early 20th century, and that there- 


ay | 


NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 273 


1:1 


fore, the two different forms might be in stable eq ium. Investigations of E. 
strigosus in Georgia document that gene flow from apomicts to sexual diploids 
may occur, such that progeny of the latter may include a significant proportion 
of apomicts (Noyes unpublished). Whether or not sexual and apomictic plants 
occur simultaneously at the same sites in Texas, and hybridize, is unknown. 
However, because the phenology of the two forms overlaps, hybridization is 
likely. Hybridization between sexual and apomictic plants has been thought 
to be important as a source of novel apomictic genotypes (Stebbins 1950) and 
has been hypothesized to be an important part of the dynamics between 
apomictic and sexual Taraxacum in Europe (Menken, Smit, @ Den Nijs 1995; 
Verduijn, van Dijk, & van Damme 2004). However, in theory, apomictic plants 
should ultimately replace sexual populations through reproductive advantage 
(Maynard Smith 1978) or by unidirectional gene flow via pollen from apomic- 
tic to sexual plants (Mogie 1992). To determine if apomictic and sexual Erigeron 
strigosus are in dynamic equilibrium in Texas, or if replacement is occurring, 
detailed demographic study will be required. 


Status of E. traversii and a new combination 

Although not known by Shinners at the time he published the species, pollen 
evidence indicates that Erigeron traversii is a sexual diploid taxon. It is there- 
fore distinct from other Erigeron strigosus in Texas, which, based on pollen, is 
apomictic and polyploid. The analysis of collection dates supports Shinners 
(1964) observation that E. traversii, on average, flowers earlier than E. strigosus. 
Although Shinners documented E. traversii in Pineywoods habitats, our data 
also document the existence of sexual diploids in Post Oak Savannah. Accord- 
ing to Shinners, E. traversii was distinct in the field from E. strigosus in produc- 
ing showier plants. Whether that difference is due to features of the capitulum 
(colors, shapes, presentation) or shape and structure of the inflorescence has 
not been evaluated. These traits, as well as geographic variation among sexual 
plants, require further study. 

The question of whether or not E. traversii should be resurrected based on 
the available evidence is not without controversy. Erigeron traversii is distinct 
from E. strigosus in ploidal level and mode of reproduction but the two taxa are 
sympatric and likely hybridize. However, the sexual taxon appears to be a con- 
sistent component of forested habitats of east Texas. To recognize the distinc- 
tiveness of E. traversii yet acknowledge the close relationship between E. 
strigosus and E. traversii, the following combination is proposed: 

Erigeron strigosus Muhl. ex Willd. var. traversii Soda roe soul & stat 
nov. Erigeron traversii Shinners, Sida 1:376. 1964. TyPE: U.S.A. T Cc 
US 59, ca. 8 mi S of Nacogdoches, 1] Apr 1942, CL. Lundelland A.AL ae eee 
BRIT). 


The holotype of E. strigosus var. traversii shows strong morphological similar- 
ity tosexual diploid plants from South Carolina, Georgia, Florida,and Alabama 


274 BRIT.ORG/SIDA 22(1) 


(Noyes & Allison 2005). In particular, these plants are distinguished by nar- 
rowly spathulate rosette and lower stem leaves that bear small, alternate, mu- 
cronate lobes along the petiole margin. Thus it is possible that E. strigosus var. 
traversii extends continuously from east Texas to South Carolina on the coastal 
plain and piedmont. Molecular phylogenetic analysis indicates that sexual dip- 
loid southeastern populations of Erigeron strigosus are distinct from sexual dip- 
loid edaphic endemic populations of E. strigosus var. calcicola and E. strigosus 
var. dolomiticola (Noyes 2006a). On the other hand, ongoing geographic map- 
ping for E. strigosus indicate that sexual diploid populations occur in western 
Louisiana, where they are contiguous with those in East Texas, and also in scat- 
tered localities in the Ozarks of western Arkansas and adjacent Oklahoma 
(Noyes 2006b). Detailed biosystematic analyses will be required to determine 
if these geographically disparate sexual populations constitute a single taxon, 
or several distinct taxa. 

The precise leaf morphology observed in sexual plants of Erigeron strigosus 
has not been observed for apomictic polyploid E. strigosus, which may entirely 
lack petiole lobes, or exhibit larger, more irregular lobing. Variability in apomic- 
tic E. strigosus is complex, extending from E. strigosus var. septentrionalis 
(Fernald & Wiegand) Fernald, which includes plants approaching E. annuus 
(L.) Pers., to apomictic Erigeron strigosus var. strigosus, which can be very simi- 
lar to E. strigosus var. traversii. In addition, the closely related species E. tenuis 
Torr. & A. Gray, also occurs in Texas and adjacent states. The likelihood of hy- 
bridization and morphological similarity between early flowering E. strigosus 
(likely corresponding to E. strigosus var. traversti) and E. tenuis was discussed 
by Van Vleet (1951). Further, based on consideration of published chromosome 
counts, F. tenuis also likely includes sexual and apomictic plants, but the geo- 
graphic distribution of mode of reproduction in the species is unknown. Thus, 
elucidation of evolutionary relationships for Erigeron strigosus must also in- 
clude detailed study of E. tenuis. 

The recognition of E. strigosus var. traversii brings to three the number of 
recognized sexual taxa related to the widespread apomictic taxa EF. strigosus, E. 
annuus, and E. tenuis and contributes to our understanding of the evolution 
and dynamics within the species complex. 

The following key is provided to aid in distinguishing Erigeron strigosus 
var. traversii from its close relatives in Texas: 


1. Pappus of ray and disc florets of numerous conspicuous capillary pappus bristles; 
the species including both sexual and apomictic plants and in need of taxonomic 
study E. tenuis 
. Pappus of ray florets a low crown-like series of scales, lacking Conspicuous capillary 
pappus bristles, disc florets with capillary bristles 
2. Plants relatively tall (to 1.5 m) with ace heal pliable leaves; dense, long, flat- 
tened trichomes present along the main stem and on the involucre; apomictic 
and usually triploid; relatively uncommon and restricted to eastern Texas E.annuus 


NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 


275 
2. Plants relatively shorter (to 0.7 m) with narrower, stiffer | flatt | trichomes 
if present, restricted to the base of the plant or relatively sparse or absent; ere 
or apomic 


tic. 
3, Plants generally lacking flattened trichomes throughout; plants sexual, dip- 
loid; pollen of high quality, relatively uniform in size, grains < 15 um diam; 
restricted to Pineywoods habitat in East Texas and adjacent Louisiana ___ E. strigosus 
var. traversii 
3, Plant stems and involucres sometimes possessing spreading flattened tri- 
chomes; plants polyploid, apomictic, highly variable in morphology; pollen of 
poor quality, highly variable in size, largest viable grains > 17 um diam; rela- 
tively widesprea 
4. Hairs cae flattened, mostly 0.5—1.2 mm; hairs of stems 0.5-1 mm 
appressed to spreadin E. stri cae var. rseptentrionali 
4, Hairs of involucres terete, mostly 0.1—-0.5 mm;hairs of stems mostly 
mm, appressed to ascending E. aa var. strigosus 


ACKNOWLEDGMENTS 


We thank the curators at BRIT and TEX for specimen loans, staff at COLO for 
providing assistance managing specimens, and Kiyoshi Yamashita, Christof 
Bigler, and Jason Knouft, of the University of Colorado, for help implementing 
GIS analyses. Guy Nesom (BRIT) and Tim Lowrey (UNM) provided valuable 
comments in review. 


REFERENCES 


Asker, S.E and L. Jertina. 1992. Apomixis in plants. CRC Press, Boca Raton, FL. 

Baker, H.G. 1965. Characteristics and modes of origin of weeds. In: G.L. Stebbins, ed. The 
genetics of colonizing species. Academic Press, New York. Pp. 147-172. 

Bark.ey, T.M.(ed.). 1986. Flora of the Great Plains. University Press of Kansas, Lawrence, Kansas. 

Beyer, H.L. 2004. Hawth’s analysis tools for ArcGIS (http://ww tialecology.com/htools). 


BierzycHubek, P. 1987. Patterns in plant parthenogenesis. In: S.C. Stearns, ed. The evolution 
of sex and its consequences. Birkhauser Verlag, Basel, Boston. Pp. 197-217 

Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner, TX. 


Cronauist, A. 1947. Revision of the North American species of Erigeron, north of Mexico. 
Brittonia 6:121-302. 

Den Nus, J.C.M.and S.B.J.Menken. 1994. Breeding systems and evolution in Taraxacum. Evol. 
Trends PI. 8:11-20. 

Grant, V. 1982. Plant speciation, 2nd. Columbia University Press, New York. 

Gustarsson, A. 1946-1947. Apomixis in higher plants. Lunds Univ. Arsskr.42—43:1-370. 

Haac, C.R. and D. Esert. 2004. A new hypothesis to explain geographic parthenogenesis. 
Ann. Zool. Fennici.41:539-544. 

Hatcu, S.L., KN. GANDHI, and L.E. Brown. 2001. A checklist of the vascular plants of Texas, 
version 2. (http://www.csdl.tamu.edu/FLORA/taes/tracy). Texas A&M University, Col- 
lege Station. 


276 BRIT.ORG/SIDA 22 


= 


1) 


Hickman, J.C. (ed.). 1993. The Jepson manual: higher plants of California. University of Cali- 
fornia Press, Berkeley. 

Hoimeren, |. 1919. Zytologische Studien uber die Fortpflzung bei den Gattung Erigeron 
und Eupatorium Kong. Svenska Vetenskapsakad. Handl. 59:1-118. 

Joxnston, M.C.1990.The vascular plants of Texas: a list up-dating the manual of the vascu- 
lar plants of Texas, 2nd.M.C. Johnston, Austin, TX. 


Kartesz,J.1.1992.A synonymized checklist of the vascular flora of the United States, Canada, 
and Greenland, 2nd. Timber Press, Portland, OR. 

LYNCH, ue 984. Destabilizing Pee general-purpose genotypes and geographic 

is. Quart. Rev. Biol. 59:257-290. 

re Smith, J. 1978. The evolution of ae University Press, Cambridge. 

Menken, $.B.J.,E.Smit,and J.C.M. Den Nus.1995.Genetic population structure in plants:gene 
flow between diploid sexual and triploid asexual dandelions (Taraxacum section 
Ruderalia). Evolution 49:1108-1118. 

Mosie,M. 1992.The evolution of asexual reproduction in plants. Chapman and Hall, London. 

Nesom, G.L. 1978.Chromosome numbers in species of Erigeron and Conyza (Compositae). 
Sida 7:375-381. 

Noyes, R.D.2006a. Intraspecific nuclear ribosomal DNA divergence and reticulation in sexual 


diploid Erigeron strigosus (Asteraceae). Amer. J. Bot. 93:470-479. 

Noyes, R.D.2006b. Reticulation and the evolution of apomixis in Erigeron sect.Phaiacroloma 
(Asteraceae) In: E. Hérandl, U. Grossniklaus, P Van Dijk, and T. Sharbel, eds. Apomixis: 
evolution, mechanisms and perspectives. Regnum Veg. 147. (Gantner Verlag, Ruggell, 
Liechtenstein) (Accepted for publication) 

Noyes, R.D.and J.R. ALuson.2005.Cytology, ovule development and pollen quality in sexual 
Erigeron strigosus (Asteraceae). Int. J. Pl. Sci. 166:149-159. 

SHinNeRS, L.H. 1964. New names and records for Texas Compositae. Sida 1:373-379. 

Soi Suavey Starr. 1999. Soil taxonomy:a basic system of soil classification for making and 
interpreting soil surveys. USDA Natural Resource Conservation Service Agriculture 
Handbook # 436, U.S. Government Printing Office, Washington DC. 

STANLEY, R.G.and H.F. Linskens. 1974. Viability tests. In: Pollen: biology, biochemistry,and man- 
agement. Springer, Berlin. Pp.67-86. 


Steesins. G.L. 1950. Variation and evolution in plants. Columbia University Press, New York. 

USDA, NRCS. 2005. The PLANTS Database, version 3.5 (http://plants.usda.gov). Data com- 
piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge, 
LA. 

Van VieeT,R.L.1951.Phenotypic variations of Erigeron strigosus Muhl.(Compositae) in east- 
ern Texas. Field & Lab. 19:161-162. 

VerRDUUN, M.H., PJ.vAN Duk, and J.M.M.van Damme. 2004. Distribution, phenology and demog- 
raphy of sympatric sexual and asexual dandelions (Taraxacum officinale s..): geographic 
parthenogenesis on a small scale. Biol. J. Linn. Soc. 82:205-218. 


MORE AMERICAN BLACK SAPOTES: NEW DIOSPYROS 
(EBENACEAE) FOR MEXICO AND CENTRAL AMERICA 


Mitchell C. Provance Andrew C. Sanders 
Depainens orp raya! and ea : ences UCR Herbarium 
College of Natural ciences Department ee and Plant Sciences 
Boies ity of Cal ifornia na University of California, Riverside 
Riverside, ee ig 92571-0124, U.S.A Riverside, ie 92521-0124, U.S.A. 
chtell@prodigy.net andrew.sanders@ucr.edu 
ABSTRACT 
During the preparation of a monograph of the Ebenaceae for Mexico, numerous llections previ- 


ously determined to be Diospyros conzattii Standley and Diospyros riojae Gomez-Pompa could not 
be reconciled with their original descriptions, type specimens and illustrations. It was determined 
that collections of putative D. conzattii from Costa Rica represent a new species, D. costaricensis, 
described here. All but one collection of putative D. riojae from the Mexican states of Queretaro, 
Hidalgo, Tamaulipas, and San Luis Potosi, represent a new species, D. gomeziorum, described here. 
Putative collections of D. fies from Veracruz additionally include specimens of D. conzattii, a spe- 


cies previously unreported for the state, and a new species from the Sierra de Los Tuxtlas, D. tuxtlensis 
described here. We provide emended descriptions of D. riojae and D. conzattii, and describe three 
id distribution, ecology 


new species of Diospyros for Tropical America. We also 

conservation, ethnobotany, and provide illustrations for peach of these taxa. 

Keyworps: black sapote, Costa Rica, Diospyros gomeziorum, D. conzattii, D. costaricensts, 
pergamentacea, D. riojae, ee rosei Complex, D. tuxtlensis, Ebenaceae, granadilla, ae 


Mexico, new species, persimmon, za gro silvestre, zapotillo 
RESUMEN 
pieen - Bispace de la ee aaa de ehaeaues para México, numerosas colecciones 
tii Standley y Diospyros I iojae Gome Pompa no 


pudieon ser reconciliadas con las Ree eons originales, el espécimen tipo y las ilustraciones. Se 
determino que la coleccion putativa D. conzattii de Costa Rica, representa una especie nueva, D. 
costaricensis descrita aqui. Todas, excepto una de las eolecone putativas de D. riojae de los estados 
mexicanos de Querétaro, Hidalgo, Tamaulipas y San Luis Poto n una especie nueva de D. 
gomeziorum, descrita aqui. Adicionalmente, las eee mietie de D. riojae de Veracruz, 
ie especimenes de D. conzattii, una especie no citada previamente para ese estado y de una 
especie nueva de la Sierra de los Tuxtlas D. tuxtlensis, descrita aqui. Proveemos descripciones 
corregidas de D. riojae y D. conzattii y describimos tres especies nuevas de Diospyros para América 
tropical. También hacemos comentarios sobre la distribucion, ecologia, conservacion, etnobotanica 


y aportamos ilustraciones de todos los taxa. 
PaLaBras CLavE: Complejo de Diospyros rosei, Costa Rica, sate panes D. conzattii, D. 
costaricensis, D. pergamentacea, D. riojae, D. tuxtlensis, Ebenaceae, especie nueva, granadilla, 


guacalillo, México, persimo, zapote negro, zapote negro silvestre 6 


SIDA 22(1): 277 - 304. 2006 


278 BRIT.ORG/SIDA 22(1) 


INTRODUCTION 


The pantropical genus Diospyros (Ebenaceae) consists of about 500 species of 
trees, shrubs, and suffrutices (White 1983). A treatment of the Mexican species 
of Diospyros in full has not appeared since Standley (1924). Modern treatments 
have dealt with smaller areas, such as Veracruz (Pacheco 1981), northeastern 
Michoacan, Guanajuato, and Queretaro (Carranza 2000), the Yucatan Penin- 
sula (Lundell 1942), and the Tehuacan-Cuicatlan Valley, Oaxaca (Kelly 2001), 
or covered portions of southern Mexico within a larger flora (Whitefoord & 
Knapp 2001). Circumscription of the Mexican and Central America taxa is not 
entirely agreed upon. However, based on our preliminary work, a reasonable 
estimate of the number of native species occurring in Mexico is around 25. 

In 1918, C. Conzatti collected a new black zapote from the coastal moun- 
tains northeast of San Pedro Pochutla, Oaxaca. He recorded some characteris- 
tics of the tree, especially those pertaining to the edible fruit, the taste of which 
he enjoyed. He believed that the fruit of the ‘zapote negro silvestre’ (Conzatti 
3167) had potential as a new tree crop (Standley 1922). This zapote was formally 
described a few years later as Diospyros conzattii Standl. (Standley 1922). Ac- 
cording to Trabut (1926), D. conzattii was introduced to California and Florida. 
However, he did not provide any specific details, and we are unaware of any 
other information that might corroborate the putative introduction of this spe- 
cies to the United States. 

Diospyros riojae Gomez-Pompa was described from one fruiting collection 
made in the Misantla region of central Veracruz, Mexico (Gomez-Pompa 1964). 
Since then, collections of putative D. riojae have been made in several more lo- 
cations in Veracruz, Iamaulipas, San Luis Potosi, Hidalgo, and numerous places 
in northeastern Queretaro. Diospyros riojae is considered an endangered spe- 
cies (Sanchez-Coello 2002 

During the preparation of a monograph of the Ebenaceae for Mexico, we 
noticed a large amount of morphological variation among collections of puta- 
tive D. riojae and D.conzattii. While we had intended to examine the taxonomy 
of both species at a later time, in view of the gered status of D. riojae,and 
asense of some urgency regarding its protection (e.g., Sosa et al. 1998), we thought 
it would be prudent to proceed with the treatment of the following taxa now, 
lest the designation of protected areas, or expensive and time-consuming eco- 
logical research, be confounded by the unrecognized presence of similar look- 
ing taxa. 

About eighty-five specimens from seventeen herbaria were examined for 
this study. With the exception of one collection from San Luis Potosi, specimens 
of putative D. riojae collected outside of Veracruz represent a new species, D. 
gomeziorum, described here. Much of the material collected in Veracruz and 
attributed to D. riojae has been found to represent D. conzattii, and a species 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSF AMERICA 279 


new to science, D. tuxtlensis, described here. Costa Rican material, previously 
ascribed to D. conzattii, clearly represents a third new species, D. costaricensis, 
which is described here. These species are best considered members of what 
we have informally recognized as the ‘Diospyros rosei Complex’ (Provance & 
Sanders 2005), a group that additionally includes: Diospyros rosei StandL., 
Diospyrosoaxacana StandL., Diospyros palmeri Eastwood, Diospyros californica 
LM. Johnston, Diospyros sonorae Standl., Diospyros texana Scheele, Diospyros 
riojae Gomez-Pompa, Diospyros conzattii Standl., Diospyros rekoi Standl., 
Diospyros torresii M.C. Provance & A.C. Sanders, Diospyros morenoi A. Pool and 
Diospyros xolocotzii Madrigal & Rzedowski. Previously (Provance & Sanders 
2005), we called into question the distinctness of D. morenoi. At that time, we 
only had the holotype to examine. Very recent collections from Chiapas, Mexico, 
have inclined us to reevaluate our position on the status of this species. We now 
believe that it is a valid taxon, and we plan to address this issue further in the 
near future. 

In this treatment we use ‘lanceolate’ in the sense of Jackson (1916), being 
broadest near the lower third of the leaf, not at mid-leaf in the sense of Stearn 
(2000). ‘Scintillant’ refers to the sparkling appearance of the epidermis of some 
plant structures when viewed under a dissecting microscope using bright light. 
Lengths for winged petioles are unavoidably approximate. Female inflores- 
cences are inferred from fruiting specimens and persistent pedicels, where they 
are otherwise unknown for these taxa. The leaf venation for most taxa is 
‘arcolanguid, defined here as an intermediate state between eucamptodromous 
and brochidodromous, in which major lateral veins nearly form distinct loops 
with superadjacent lateral veins, but tardily wane, becoming difficult to dis- 
tinguish as loops. Tne term is coblas for Simtel ley and out of necessity, since 
the term ‘ lod ’ of Todzia and Keating (1991) 
refers toa different type of pone Geographical coordinates, elevations, 
and similar estimations when made by us are presented in brackets. 


TAXONOMIC TREATMENT 
1. Diospyros riojae Gomez-Pompa, J. Arnold Arbor. 45:465. 1964. (Fig. 1). Type: 


MEXICO. VERACRUZ: between Chiconquiaco and Misantla, in Liquidambar-Quercus forest, 
with Magnolia, Meliosma, Juglans, and Tu ae nia, 1350 m, 13 Aug 1962 (fr), A.Gémez-Pompa 
789 (HOLOTYPE: A; ISOTYPES: MEXU, US on fiche from UCSBI). Original material examined: 
(TYP. 1C.: A.Géomez-Pompa 7891, J. Arnold hee 45:467. 1964.). 


Trees, rarely shrubs, probably facultatively deciduous, to 25m tall,and to 2.5m 
in diameter. (Pacheco 1981); trunk channeled, rimose, bark dark-brown (Pacheco 
1981); aged stems subterete to terete, bark shallow-fissured with dark ridges, 
gray and black, stemwood pale yellow; 2nd-3rd year stems short-fissured, gla- 
brous to furfuraceous pulver ulent, punctic ulate, lenticellate, half-netted, gray 
above, tawny below; current year’s stems angular, smooth to sulcate, glabrous 


BRIT.ORG/SIDA 22(1) 


Fic. 1. A-G. Diospyros riojae Gomez-Pompa. A. Abaxial surface (left) and adaxial surf ight) surface of staminat 
petal lobe. B. Adaxi Mt ft) and | ralifig) ) f staminate infl bract. C Tpicaltanitier D. Staminate 
flower. E. Abaxial f f sh il of F. Fruiting stem. G. Fruit. A—D. 


Based on F Ventura A. 15078. E-F. Based on J. RecorraiZ: 83.G. Based ane Pacheco & J.I. Calzada 18 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 281 


to sparingly subappressed puberulent, hirtellous at the base and near the apex, 
tan to reddish-brown, glandular. Petioles stout, thicker than width, (2-)4-6(-7) 
mm long, winged, rugose below, slightly pruinose, deeply concave to V-grooved 
above, ciliolate, glandular. Leaves alternate, simple, entire, subcoriaceous to co- 
riaceous, (5-)7-13(-14.2) cm long, (2.5-)3-6(-7.5) cm wide, elliptic, widely el- 
liptic, oblong, widely obovate, sub-oval, apex acutely to obtusely rounded, or 
short-acuminate to a rounded point, margin slightly recurved, + thickened, 
sometimes ciliolate, base obtusely rounded, cuneate, or short-acuminate; lower 
leaf surface glabrous, puncticulate, sometimes slightly pruinose or scintillant, 
shiny, olive to bronze to brown; upper leaf surface glabrous, shiny, pale light- 
green to gray-green or copper-green, epidermal cells enlarged, the anticlinal 
walls thickened. Venation arcolanguid; midrib prominent below, chartreuse to 
reddish-brown, shallowly concave above, smooth to clavate glandular hairy, 
shiny, olive to yellow-green; 2°-3° veins reticulated, raised below, fine with sharp 
relief, sometimes pruinose-scintillant, granular-papillose, shiny, usually promi- 


nent above, lateral veins 8-12 pairs. Laminar extrafloral nectaries abaxial, near 
base and midrib, round to narrowly elliptic, sometimes on lateral veins. Male 
inflorescence fascicle, 1-4 flowers from leaf axils of the previous year’s growth, 
or from the base of new stems, sometimes a pair of solitary flowers from the 
new stem just above the fascicle; pedicels +-7(-10) mm long, sparingly brown 
to claret velutinous, epidermis green; pedicel bracteoles 2, | mm long, oblong, + 
flat to slightly concave, ascending wavy hairy below, the hairs thick, glisten- 
ing. Male flowers 5-merous; calyx infundibuliform; calyx tube 1.5 mm long, gla- 
brous or scantly minute subulate hairy; sepals triangular to ovate, 1.3-1.8 mm 
long, 0.7-0.9 mm wide, ciliolate, scantly minute appressed subulate hairy ba- 
sally, becoming dark glandular wavy hairy apically; corolla urceolate-campanu- 
late, tan to brown, reportedly white in living material; corolla tube constricted 
below the lobes, 4 mm long, 3 mm wide, interior grading from glabrous dis- 
tally to minutely deltoid scaly at mid-tube, again becoming glabrous near the 
base, except near filament bases, exterior puberulent with hairs erratically di- 
rected, some glandular; corolla lobes oblong-squarish, emarginate, 2 mm long, 
2mm wide, interior pubescent, exterior minutely sericeous-puberulent, 
exmedially appressed, several black hairs present near the apex. Stamens 18, 
subexserted, adnate to corolla at mid-tube or lower, some inserted on the recep- 
tacle, some lower stamens extrorse, abruptly geniculate, and connected to introrse 
upper stamens by a superdermal vascular trace; filaments | mm long, hirtellous, 
especially on margin; anthers basifixed, lance-ovate, + 2.5 mm long, laterally 
concave, apically constricted, opening by short apical slits; pistillode + wheel- 
shaped, lobes 5, bifurcate, glabrous, dark gray. Female inflorescence flowers soli- 
tary, from caducous bract scar axils near the base of young stems. Fruiting pedicels 
7-12mm long, stout, very minutely hirtellous. Fruiting calyx 5-lobed; fruiting calyx 
tube 3.5-4 mm long, explanate to ref lexed; fruiting sepals accrescent, coriaceous, 


282 BRIT.ORG/SIDA 22 


— 


) 


arcuate-reflexed to spreading, 7-25 mm long, 7-13 mm wide, oblong, narrowly 
triangular or ovate, apex usually ascending to incurved, sometimes glaucous- 
pruinose to scintillant. Fruit berry, 3-5.5cm long, up to 4. 7cm mace, SHE Opes, 
locules8-10; mesocarp [leshy, yellow-orange in dried 

and I epidermis smooth to bullate, shiny in living s material, oreen to brown 
or black, herbarium material reddish-brown to dark-brown, immature fruits 
sometimes atropurpureous. Seeds not seen in available material (seeds orangish 
in photograph in Sanchez-Coello 2002 from Conde 2000), described by Pacheco 
(1981) as brownish to grayish, shiny, very rugose, and with a prominent raphe. 


Additional specimens examined. MEXICO. oa ae Potosi. Mpio. Tamazunchale: Tamazunchale, 
[21° 16'N, 98° 47'W, 153 ml, 24 Jul 1937, M.T.E ), US). Veracruz. Mpio. Alto Lucero: Cruz 
Blanca, 19° 46'N, 96° 40'W, 900 m, 30 Apr 1980, L. Pacheco & J.1. Calzada 23 & 24(XAL); same locality 
19 Sep 1979, L. Pacheco & J.I. Calzada 18 (X AL); Cruz Blanca, 19° 40'N, 96° 50'W, [900 ml], 25 May 1998, 
Tono Vazquez 98-045 (UCR). Mpio. Chiconquiaco: Canada del Huerfano, 19° 49'N, 96° 48'W, [900- 
1200 ml, 10 Jul 1966, A. Gomez- ie 1602 (XAL). Mpio. Martinez de la Torre: Malaupan [Santa Ana 
Maloapan, 20° 04'N, 97° 04'W], LOO m, 15 Mar 1978, F. Ve He 15078 (EB, XAL). Mpio. Tantima: 
Sierra de Tantima, 21° 17'N, 97° 51 7: 740 m, 23 Aug 1979, .1. Calzada 5550 (XAL). ae Tepetzintla: 
Sierra de la Pena Blanca, Sierra de Tantima, [21° 13'N, 97° 55'W, = 700 ml, 21 Sep 1989, P Zamora C. et 
al. aes ), San José de Copaltitla al NE de Tepetzintla, 21° 12'N, 97° 52'W, 350 m, 28 Aug 1981, G. 
Castillo C.& A. Benavides M. 2265(XAL ; de Mixtepec, Sierra de Tantima, 21° 18'N, 97° 50'W, 
LOOO m, ; va 1988, J. Becerra Z.83(XA 


Distribution and Ecology.—The a known collection of this taxon made out- 
side of the state of Veracruz is from Tamazunchale, San Luis Potosi. This 1937 
collection (M.T. Edwards 670) is the earliest one of the species known to us. 
Tamazunchale is at 153 m, a low elevation based on the literature (e.g. Pacheco 
1981). In northern Veracruz, this species occurs in the Sierra de Tantima. In cen- 
tral Veracruz, occurrences are in the Sierra de Chiconquiaco, the lowlands of its 
northern base, and the Misantla region. Populations in northern Veracruz are 
in isolated cloud forest patches. In central Veracruz, collection sites have been 
characterized as forested escarpments near pasture (Sosa et al. 1998), cloud for- 
est, and deciduous forest (Pacheco 1981) at 750-1000 m. It seems not to be widely 
realized that this taxon also occurs at lower elevations at 100-300 m. Low el- 
evation occurrences are associated with tropical evergreen forest with Quercus. 
In this community, they are known to flower during March. This species is con- 
sidered to be in danger of extinction (Sanchez-Coello 2002). 

Ethnobotany.—The tree is called ‘granadilla’ in the Sierra de Tantima re- 
gion of northern Veracruz where it is considered edible (G. Castillo C. & A. 
Benavides M. 2265). The name '‘sapote prieto’, is indicated on the collection from 
Tamazunchale, San Luis Potosi. 


2. Diospyros gomeziorum M.C. Provance & A.C. Sanders, sp. nov. (Fig. 2e). Typr: 
MEXICO. QUERETARO: Mpio. Jalpan de Serra, 5-6 km al NW de San Juan de ee Duran, El Ar- 
royo, 21.48°N, 99.12°W, escaso, bosque de pino-encino, cedro blanco, orilla de arroyo, canada 

1500-1600 m, 1 Aug 1991 (fr), Benito Servin 1274 (HOLOTYPE: IEB-150586). 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 


Fic. 2.A—-E. Di ji M.C. Provance & A.C. Sanders, sp.nov. A.5 


fl B. Typical anther. C. Adaxial 


¢ L 


surface of staminate petal lobe. D. Lateral v 


E Fruiting pene A-C. Based on R. 


Ferndndez N. 2829. D. Based on Hiram Rubio 1520. E. Based on Benito Servin 1274. 


284 BRIT.ORG/SIDA 22(1) 


Arbor [rutexve usque ad 20 m altus, D. riojae Gomez-Pompa similis sed differt margine foliia pagina 


je 


adaxiali usque ad paginam abaxialem exter obliquantibus; cellulis epidermalibus paginae laminae 
= ed oe 16 tie cl 7 ; ; i} Vaal peed il 1 Ee Be 
. LS [ 
] : : ] ie sf; igs sai fb ~ ] | ot \; 14 
[ C rT i t t 7 
frouctiteris 7 


] 


Trees or shrubs to 20 m tall, facultatively deciduous; trunk reported to 12 cm 
diameter for a 5-6 m tree (Servin 1274) and 20 cm in diameter for a 4+ m tree (R. 
Fernandez N. 2459), erect, longitudinally fissured, bark gray, reportedly shiny; 
aged stems angular, bark verrucose, scarred, short-fissured, cinereous, stem wood 
yellow to yellow-orange; 2nd-3rd year stems angular to subterete, sometimes 
hirtellous, + half-netted, the outer epidermal layer gray, tawny or reddish- 
brown, the lower layer cream to orangish; lenticels protruding, sometimes 2-3 
mm long; current year’s stems subterete, smooth to striate, densely hirtellous, 
reddish-brown, puncticulate. Petioles stout, often thicker than wide, 2-5(-8) 
mm long, glabrous to minutely erect hairy to deltoid scaly, rounded to rumi- 
nate below, usually glistening, dark brown to olive green, narrowly channeled, 
flat or barely raised, furfuraceous-glandular, ascending ciliolate on margin, yel- 
low-green to olive above. Leaves alternate, simple, entire, pergamentaceous to 
chartaceous, sometimes subcoriaceous, lanceolate to ovate to elliptic, 6-11(-1L5) 
cm long, 2-4.8 cm wide, apex acute, obtuse or acuminate to a rounded point, 
margin beveled outward from upper surface at + 45° to the lower surface, some- 
what sharp, shiny, sparsely to moderately ciliolate, base acute to obtuse, decur- 
rent onto petiole; lower leaf surface glabrous to sparsely subappressed pubes- 
cent, hairs black or reddish, most common near the base, midrib, and apex, 
surface shiny, green to olive, puncticulate; upper leaf surface glabrous, glau- 
cous to pruinose-scintillant, localized or covering most of surface, but not on 
the beveled margin, granular-papillose, olive or pale blue-olive to dark-olive or 
dark blue-green, usually darker above than below, young leaves tend to be lighter 
above. Venation arcolanguid, granular-papillose, glaucous to pruinose- 
scintillant; midrib prominent below, glabrous to subappressed pubescent, hairs 


— 


— 


dlack, reddish-brown, or white, narrowly canaliculate above, sparsely to densely 
hirtellous to deltoid scaly, rarely glabrous, yellow-green; 2° venation 
subprominent below, lateral veins (8-)11-13(-14) pairs; 3° venation reticulate, 
obscure to prominent below, usually prominent above. Laminar extrafloral nec- 
taries abaxial, minute, circular. Male inflorescence fascicles of 1-3(-5) flowers 
in leaf scar axils of previous year’s growth, or one to several flowers at the base 
of new growth in scar axils of caducous bracts; pedicels 1-3 mm long, densely 
straight hairy, the hairs reddish, black, and white; pedicel bracts 2, 2-2.5 mm 
long, + linear. Male flowers (4-)5(-6)-merous; calyx infundibuliform-campanu- 
late, deeply lobed, interior glabrate or scantily minute white hairy, exterior 
sparsely minute deltoid scaly; calyx tube |.5-2 mm long, 2.5-3 mm wide, sparsely 
subappressed puberulent; sepals lance-ovate to triangular, 2-7 mm long, l-4 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 285 


mm wide, exterior glabrous to pubescent basally, subappressed pubescent 
apically, the hairs glossy, black to reddish, simple and glandular, epidermis vis- 
cous; corolla urceolate-campanulate, white in living material; corolla tube 3- 
5.5 mm long, 3-6 mm wide, exterior densely pubescent, most hairs distally ori- 
ented, some hairs irregular, interior sparingly hirtellous, hairs sometimes 
erratically directed; corolla lobes (4-)5, spreading, oblate to squarish or widely 
obovate, often emarginate, 2-4 mm long, 2-3.5 mm wide, several short black 
hairs near the apex, interior sericeous. Stamens 20, equal to, or barely surpass- 
ing tube, most adnate to corolla, some inserted on the receptacle; filaments 0.3- 
0.8 mm long, ascending puberulent; anthers basifixed, oblong-ovate to ovate, 2 
mm long, opening by short slits near apex, ay by cone: lateral ne 
granuliferous, apex acute or rounded an 
wheel-shaped, short, wide, corrugate. Female irifloreseenee flowers eee 
cauline near the base of current season's growth, rarely from leaf scar axil of 
old growth (e.g. Ferguson 15), sometimes two opposite inflorescences, but two 
mature opposite fruit not seen. Female flowers not seen, but styles (4-)5 as rem- 
nants on mature fruit, appressed white puberulent. Fruiting pedicels (4-)5-8(- 
11) mm long, densely hirtellous. Fruiting calyx 5-6-lobed. Fruiting calyx tube 3- 
4 mm long; fruiting sepals accrescent, + spreading, 19-20 mm long, 8-11 mm 
wide, lanceolate to narrowly elliptic or obovate, glabrous to sparsely appressed 
hairy, apices rounded, sometimes ascending, surfaces often bluish-white glau- 
cous, non-glaucous areas tawny, puncticulate. Fruit berry, subglobose to slightly 
obovoid, 3cm long, 3-4.5 cm in diameter, green when immature, turning black 
(and then presumably ripe), atropurpureous in dried specimens, locules 10; 
mesocarp orange when dry; hypodermis thin, not very stony, often broadly 
rippled in specimens; epidermis orange-peel-textured, scintillant, glaucous- 
pruinose. Seeds 16-19 mm ee 9-10 mm wide, rugose, shiny, light brown. 


Paratypes: MEXICO. Hidalgo. Mpi B le Meztitlan, 9 km al NE de Mesquititlan a lo 
largo de una canada, 20.74°N, 98. 98° W, 2200 m, 3 May 1975, FG. Medrano et al. 7932 (SD, US, XAL). 
Queretaro. Mpio Arroyo Seco: 2 km al W de El Jardin, 21° 25' 24"N, 99° 41' 42"W, 1380 m, 4 Jul 1989, F. 
Carranza 1852 (1EB). Mpio. Jalpan de Serra: 2-3 km al Poniente de San Isidrio, La Parada, 21° 30.5'N, 
99° 10'W, 1300 m, 14 May 1990, Benito Servin 206 (CIIDIR, IEB); 2-3 km al S de La Parada, 21° 32.3'N, 
99° 10'W, 1200 m, 26 Mar 1990, Benito Servin 58 IEB); 6-7 km al Oriente de La Parada, 21° 30.5'N, 99° 
6.8'W, 1400 m, 5 Apr 1990, Benito Servin 96 (EB); 2-3 km al N de La Parada, 21° 32.3'N, 99° 10'W, 1100 
m, 13 Aug 1990, Benito Servin 412 (EB); + 2 km al W de La Parada, Cuesta de los Lirios, 21° 30.5'N, 99° 
11.2'W, 1180-1250 m, 13 Mar 1990, E. Carranza 2392 (C oie IEB). Mpio. Colon: cerca vado rio Colon, 
20° 48'N, 100° 03'W, oe m, 2 Apr 1982, Elizabeth Arquti 1761 ae ies Pinal de Amoles: | km 
S de Escanelilla, sobre la carretera a Pinal de ee 21° ON, e 33'W, 1250 m, 18 May 1987, J. 
Rzedowshi 43398 (HUAA, IEB); + 4 km al SE de Santa Agueda, 21° 14' 30" N. 99° 37'54"W, 1190 m, = 
Apr 1989, E. Carranza 1635 (IEB); 3 km al S de Escanelilla, 21° 10.2'N, a 34'W, 1100 m, 19 Mar 1985, 

Ferndndez N. 2829 (US); same location, 30 Jul 1984, R. Fernandez N. 2459 (LEB). Mpio. Landa 

Matamoros: |.5 km al SW de El Naranjo, 21° Ol 18"N, 99° 27' 42"W, 900 m, 10 Mar 1990, Hiram a 
1542 (IEB); 1 km al Poniente de El Cerro de La Palma, 21° 12' 28"N, 99° 04' 32"W, 1500 m, 2 Mar 1990, 
Hiram Rubio 1520 (IEB); cerca de Tres Lagunas, 21° 19' 36"N, 99° 12' 12"W, 1700 m, 22 Jun 1988, J. 


286 BRIT.ORG/SIDA 22(1 


Rzedowski 46695 (IEB); 2 km al Norte de Neblinas, 21° 13' 58"N, 99° 06'18"W, 1060 m, 29 Sep 1988, 
Hiram Rubio 185 (EB); El Humo, 2 km a TOuest d’El Humo, 21° 18'N, 99° 05'W, 1320 m, | Oct 1994, J.N. 
Labat & FE. Carranza 2562 (EB); 2 km al SE de El Humo, 21° 13'55"N, 99° 06'15"W, 1100 m, 7 Jun 1989, 
Hiram Rubio 775 (HUAA, IEB); 2.5 km NE de El Humo, [21° 18'N, 99° 05'W], 1000 m, 11 Aug 1990, 
Hiram Rubio 1865 (EB); 1.5 km SE de El Naranjo, 21° Ol' 18"N, 99° 27' 42"W, 1000 m, 25 Jun 1990, 
Hiram Rubio 1746 (EB), 4-5 km al S de Tres Lagunas, 21° 19’ 33"N, 99° 12'12"W, 1820 m, 3 Mar 1990, E. 
Carranzd 2359 (HUAA, IEB), El Calvario, | km al Poniente de El Aguacate, 21° 16' 06", 99° 14' 42", 1620 
m, 26 Jun 1989, Hiram ae 838 (EB); + 8 km de desviacion, camino Tres Lagunas, 21° 19' 35"N, 99° 
06'15"W, 1900 m, 7 Nov 1988, E. Carranza 1144 (EB); 2 km al SE de Neblinas, 21° 15'N, 99° 03'12"W, 
900 m, 23 Jun 1989, Hiram Rubio 826 (EB). San Luis Potosi. Mpio. Rayon: 3.5 mi by road Sof km 81.5 
onhwy 70 Wol Valles at microwave tower,[+ 21° 53'N, 99° 30'W], 1340 m, 23 May 1981, George Fergu- 
son 15 (UTEP). eae Mpio. Gomez Farias: Rancho del Cielo, a 11 km de Gomez Farias, [23° 
O4'N, 99° 12'W], 1110 m, 22 Sep 1974, EG. Medrano 7416(ARIZ); Rancho del Cielo, La Sierra de Gomez 
Farias, 1100 m, 1 May 1982, H. Narave F. & T. Dent 134 (XAL); Rancho El Cielo, F CATR E 
ridge of the Sierra Cucharas, 1200 m, 23 Aug 1950, E. Hernandez X.& F. Harrison X-586] (CHAPA 


Distribution and Ecology.—This specie urs in the Sierra Madre Oriental, in- 
cluding the Sierra Cucharas, et north of Gomez Farias in southwest 
Tamaulipas, t thwest of Tamasopo in southeast San Luis 


Potosi, northeast Queretaro and Barranca de Meztitlan in eastern Hidalgo. Col- 
lections are from a variety of vegetation types at elevations between 900 and 
2200 m. More material, by far, has been collected in the state of Queretaro than 
any other state. In Queretaro, it has been come as abundant in pine-oak 
forest, oak forest, oak forest with Liquid and Tilia forests. Other settings 
in which it has been reported but deserved. as scarce include gallery forests 
with Platanus, and canyons with bosque mesofilo de montana (+ cloud forest). 
An unusual association occurs in Hidalgo, where it was described as very abun- 
dant in a canyon with matorral espinoso and matorral crasicaule at 2200 m 
(EG. Medrano et al. 7932). The five male flowering collections we examined were 
all taken in March. Some of these specimens were just leafing out at the time. 
The fruits ripen between September and early November. 

Ethnobotany.—Queretaro: ‘zapote prieto’ (Rubio 1746), ‘zapote del monte’ 
(Servin 206), zapotillo (Carranza 2000). 

Etymology.—The epithet is intended to honor both Arturo Gomez-Pompa, 
who discovered and described Diospyros riojae, and has also been a great inspi- 
ration to our work on Latin American Diospyros, and also the late J. ‘Carmelo’ 
Gomez, who assisted the first author in the field on many occasions, and was 
very knowledgeable on local plant use in the Sierra Tlachichila, Zacatecas. 

Collections of D. gomeziorum have often been confused with D. riojae. A 
conspicuous feature that differentiates D.gomeziorum from D. riojae is the leaf 
margin, which is beveled outward from the upper to the lower surface at about 
45° in D.gomeziorum. The margin formed tends to be sharp, shinier, and greener, 
while that in D. riojae is thick, not as shiny, and a paler green. Other differences 
include the upper leaf surface, which is often copiously glaucous-pruinose in 
D. gomeziorum, while typically only the abaxial leaf surface is slightly prui- 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 287 


nose-scintillant in D. riojae. The epidermal cells of the upper leaf surface in D. 
riojae are large, with thickened anticlinal walls, a feature absent in D. 
gomeziorum. Leaf shape in D. gomeziorum ranges from elliptical to lance-ovate 
or ovate, while the leaves of D. riojae often tend to be oblong or obovate. How- 
ever, this character overlaps, since both taxa may have elliptical leaves. In fruit- 
ing collections, the shorter, less stout, fruiting pedicels, the thinner, less scleren- 
chymatous fruit wall, and the glaucous-pruinose to scintillant fruit distinguish 
D. gomeziorum. The pedicel bracteoles of the male inflorescence in D. 
gomeziorum are twice as long as those in D. riojae, and are nearly folded length- 
wise (navicular). The anthers in D. gomeziorumare minutely granular, lacking 
the apical constriction and smooth texture seen in D. riojae. 

Carranza (2000) described male flowers of this species (as D. riojae) as cy- 
mose. We did not find cymes in material referable to either species. Develop- 
mentally, the inflorescences may be related to cymes, but, they appear to be 
fascicles. Some flowers may appear to be in dense terminal clusters (e.g., H. Rubio 
1542). However, this appearance is superficial, since small terminal shoot api- 
ces can be found, although sometimes only with difficulty. 


3. Diospyros conzattii Standl., J. Wash. Acad. Sci. 1217):399. 1922. (Fig. 3). Tyee: 
MEXICO. OAXACA: Distrito de Pochutla, Cerro Espino, Cafetal oa ead a a 1917 (fr), C. 


Conzatti 3167 (HOLOTYPE: US-1014759 not seen; ISOTYPES: MO-8 6! Gi 1100 m, with 
Reko & Makrinus), MO-1039787!, fragment US-892600! (in ene 900 m, Gch Reko & 
Makrinus). 


Trees or shrubs, 8-10 m tall, probably facultatively deciduous, trunk not re- 
ported; aged stems subterete to angular, bark verrucose, fissured, gray, stemwood 
off-white to yellowish or pale orange; 2nd—-3rd year stems angular to subterete, 
bark rimose, epidermis glabrous to sparsely hirtellous, sometimes pulverulent, 
sparingly clavate glandular hairy, densely lenticellate, puncticulate, chestnut 
to tawny, becoming silvery gray; current year’s stems quadrangular, finely sul- 
cate, glabrate to hirtellous, sometimes sparsely strigillose, the hairs off-white 
to brown, the epidermis shiny, viscous, olive to nearly black. Petioles usually 
thin, sometimes flexuous, 5-8 mm long, pale green to green-brown, rounded 
below, glabrous to appressed puberulent, conspicuously winged part way above, 
the wings becoming vertically oriented and narrow along the petiole, petiole 
slightly convex and minutely V-grooved above, to widely flat-channeled, to 3- 
channeled (main channel raised and skirted by side channels formed by verti- 
cal wings), glabrous to erect puberulent, sometimes clavate glandular hairy, 
especially in young leaves. Leaves alternate, simple, entire, pergamentaceous to 
chartaceous, lanceolate to elliptic to ovate, 5-14(-14.7) cm long, (2.5-)3-5(-6) 
cm wide, immature leaves membranaceous, apex acuminate to an acutely or 
obtusely rounded point, margins flat to subrevolute, curved downward near the 
base of the leaf, thickened intramarginal zone seen when viewed abaxially, 
hyaline and minutely ciliolate in immature leaves, base acutely or obtusely 


288 BRIT.ORG/SIDA 22(1) 


di A Fruiti } hlet RS H £] . L h C. Adayial £ £ ct 


GPEA CA tr apu 


Fic.3.A 
nate petal lobe (eft) ant Hplal anther (ight). D.S 


41.] + eee ten anaes aac edon 


JL. Matinee A, Hernandez 1302. B- E. Based on F. Ventura A. 7131. 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 289 


rounded, someti te, decurrent onto petiole, sometimes loosely recurved 
near the petiole; lower leaf surface usually glabrous, very rarely minutely white 
glandular hairy, puncticulate, dull, olive; upper leaf surface glabrous, glaucous, 
densely puncticulate, dull, olive. Venation arcolanguid; midrib very prominent 
below, slightly undercut, terete, glabrous, straw to light-green, weakly raised 
above, and then caniculate within the raised midrib, very sparsely minute erect 
hairy; 2°-3° venation reticulate, veins narrow, raised below and above, lateral 
veins 9-14 pairs, 2° arches formed with superadjacent lateral veins typically 
obscure. Laminar extrafloral ies abaxial, conspicuous, round to oval, dark- 
red to black, often evenly spaced and near the midrib. Male inflorescences cymes, 
I(-2) near the base of current year’s stems, or (1-)2 leaf scar axils of the previ- 
ous year’s growth, 1-3-flowered, black to umber velutinous, sparsely covered 
with minute clavate glandular hairs; peduncles 4-6 mm long, velutinous, um- 
ber to dark brown; pedicels slender, 5-7 mm long, vestiture as in peduncles; 
pedicel bracts 1-2, alternate or opposite, narrowly oblong, 1 mm long, 0.8 mm 
wide, umber. Male flowers 5(-6)-merous; male flowering calyx funnelform, dry- 
ing very dark-brown to black, glabrate to appressed-pubescent, coterminous 
with pedicel and uninterrupted; male flowering calyx tube 2-3 mm long, 1.5-2 
mm wide; sepals 5, acute-triangular, 2-4 mm long, ciliolate, apically vermiform 
glandular hairy, sinuses rounded; corolla 5-6-lobed, long-urceolate, drying very 
dark-brown to black, reportedly whitish in life; corolla tube widest above middle, 
7-8.5 mm long, 4-5 mm wide, abruptly constricted distally, exterior densely 
minute white puberulent, hair density + increasing distally, interior sparsely 
white puberulent, the hairs concentrated in regions of filament attachment; 
corolla lobes quadrate to oval, 2-2.5 mm long, 1.5-2 mm wide, asymmetrical, 
apex obtuse-rounded, truncate, emarginate, or trifid, often with several short 
dark hairs, exterior densely minute white puberulent, interior inflexed at the 
distal right margin, left margin slightly involute, pubescent. Stamens 30, some- 
times attached in pairs, adnate to corolla from midpoint to bottom of tube, some- 
times inserted on the receptacle; filaments 3 mm long, minutely hairy; anthers 
basifixed, lanceo-apiculate, 3-3.5 mm long, opening by terminal pores or short 
lateral slits, connective minutely deltoid-scaly, pistillode minute, obturbinate, 
smooth, nearly unlobed, having a few smooth very small basal lobes, glabrous. 
Female inflorescence {lowers solitary, cauline, at the junction of current and 
previous year’s growth. Fruiting pedicels stout, (7-)12-15 mm long, glabrous to 
sparingly minute hirtellous. Fruiting calyx 5-lobed; fruiting calyx tube 3 mm 
long, explanate to ee usually ae a conspicuous enlarged basal re- 
gion encircling the pedicel joint; fruiti t spreading to reflexed 
slightly, apices straight, coriaceous, narrowly to linearly triangular, 24-3 Lin 
long, 5-8 mm wide, margins sometimes sharp edged, sometimes glaucous, gla- 
brous, puncticulate. Fruit berry, depressed-globose slightly obovoid, up to 4cm 
long, 4.5 cm in diameter, reportedly reddish (Chazaro 3969) or green (Standley 


290 BRIT.ORG/SIDA 22(1) 


1922) when ripe, mostly atropurpureous in herbarium material. Locules 10-12; 
mesocarp reportedly black when ripe, usually orange in dried specimens, hy- 
podermis 0.2-0.5 mm thick, stony; epidermis bullate, glaucous-scintillant, usu- 
ally wrinkled when dry. Seeds 15-16 mm long, 11-12 mm wide, 5.5-9.5 mm thick, 
sculpture cerebriform, chestnut. 
Additional material examined: MEXICO. Veracruz. Mpio. Calcahualco: 4.2 km W of Escola, 19° LO'N, 
97° 10'W, 2200 m, 12 Jan 1981 (fr), M. Nee & G. Schatz 19777 (XAL). Mpio. Chiconquiaco: abajo de 
el reuvane Arroyo Colorado, 19° 46'N, 96° 45'W, 1650 m, 25 Oct 1988 (fr), C. Gutierrez B. 3292 
XAL). Mpio Coscomatepec: 3 km al SE de la antigua Xicola, Cima del Cerro La 
ee i; O6'N, 97° 04'W, 1900 m, 29 Apr 1987 (fr), J.L. Martinez & A. Hernandez 1302 (XAL). 
Mpio. Jalacingo: El Cuizalin, ca. 19° 50'N, 97° 16'W!, 1500? m, 22 May 1982 (pist. fl), F Ventura A.7131 
(IEB, MO, XAL). Mpio. Tatatila: camino de herradura de Tatatila a Escalone (o Puente Caballos), [+ 
19° 42'N, 97° 6'W, 1500 mJ, 14 Jan 1986 (fr), M. Chazaro & Roberto Acosta 3969 (WIS, XAL). Mpio. 
Tlacolulan: Abajo del Saucal, direccion Agustin ie 19° 45'N, 96° 57'W, 1180 m, 14 Aug 1990 (fr), 
C. Gutierrez B. 4056 (XAL). 
Distribution and Ecology—The type locality is in the Pacific coastal ranges of 
the Sierra Madre del Sur, Oaxaca. It occurs there in dry tropical forest on coastal 
slopes (Anonymous 1927). According to Conzatti (in Standley 1922), the fruit is 
ripe in April. In Veracruz, occurrences are near Pico de Orizaba, the Sierra de 
Tezuitalan, and the Sierra de Chiconquiaco, between 920 and 2200 m altitude. 
It occurs on slopes with pine-oak woodland and Alnus, oak forests, and decidu- 
ous forests. 

Ethnobotany.— Veracruz: ‘zapotillo’ (C. Gutierrez B. 3292). Oaxaca: ‘zapote 
negro montés’ (Standley 1922 [quoting Conzattil, and on US-892600), ‘zapote 
negro silvestre’ (Conzatti 3167 [in his own handwriting). The wood is consid- 
ered valuable (Anonymous 1927 [paraphrasing Conzatti)). 

Only one of the isotypes we examined, MO-879066, included a fruit. Un- 
fortunately, the ges atropurpureous fruit isin rather poor condition. However, 

on close examination it was clear that the epidermis is bullate, glaucous, al- 

though ae towards brown in some areas. The fruit wall is thick and quite 
stony. A fragment packet attached to another type, MO-1039787, contains a 
spreading fruiting calyx tube bearing the basal 5 mm of one sepal, and much 
less of the base of another. The sepals appear to have been quite narrow, ca. 4-5 
mm wide at the base and further narrowed distally. Characteristics seen in the 
fruit of D. conzattii collections from Veracruz are consistent with the charac- 
teristics found in the isotype. An enlarged region below the fruiting calyx en- 
circling the pedicel joint is only seen in the Veracruz material, but not the 
isotype. At this point, we are unsure of its significance. 


'Coordinates based on F. Ventura A. collection of Rhamnus capreaefolia var. capreaefolia from Cuizalin, 22 June 
1970 (LL) accessed through REMIB. www.conabio.gob.mx/remib_ingles/doctos/ remib_ing.html. 


The label indicates the collection was made at 150 m. This seems to be a typ he . The coordinates 


f isali thi th Hi + 
for Cuizalin p | 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 291 


4. Diospyros costaricensis M.C. Provance & A.C. Sanders, sp. nov. (Fig. 4b, e). Tyre: 
ICA. GUANACASTE. Canton de Liberia: Parque Nacional de Guanacaste, Cordillera 
de Guanacaste, Cerro Cacao, Estacion Cacao, 10° 55' 45"N, 85° 28' 15"W, 1100 m, 14 Jul 1991 (fr), 
Carlos Chavez 569 (HOLOTYPE: MO-5316680!; IsoTyPEs: “4 duplicates” indicated on label, but 
not seen by us). 


poet D ttii Stand. similis sed ides-faciente, usque ad 35 m alta et 1.5m diametro; petiolis 


lil neque elevata in Baend abana 


1 
] ( ] \ 


costa cao ad erroniel adaiale, | 


longictroreim cricnantiin ti 


pk 


Trees, probably pelea dy deciduous, reportedly colonial, reportedly up to 
35 m tall, and 15 m in diameter (Espinoza 54); trunk buttress-forming, chan- 
neled, smooth, greenish-black to greenish-brown, slash yellow and aromatic; 
aged stems somewhat angular, half-netted, black to gray above, lower layer beige 
to golden-brown, sometimes + mottled, sometimes shallow-fissured, 
lenticellate; 2nd-3rd year stems irregularly half-netted, beige beneath, black 
and grey above, hirtellous, pubescence ee persisting on three year old 
stems, lenticellate; current year’s st sulcate to minutely ridged, 
sparsely subappressed Bolden hairy, the hairs fine and = straight, sparingly to 
very densely erect puberulent, dark-green to black, somewhat glandular, be- 
coming lenticellate. Petioles usually thin, somewhat flexuous, 3-6.5 mm long, 
wings tapering gradually from the decurrent lamina, and twisting abruptly 
into a vertical orientation along the margin, glabrous to hirtellous below, rug- 
ose to invaginate, light olive to very dark brown, concave above, often minutely 
V-grooved, usually golden hirtellous, often glandular, epidermis often nearly 
black. Leaves alternate, simple, entire, chartaceous to subcoriaceous, cartilagi- 
nous, ovate to elliptic, rarely widely-obovate, 6.5-9.5(-10.2) cm long, 2.8-4(-5) 
cm wide, very often tattered around the margins, wrinkled, apex usually asym- 
metric, tapering to an obtusely rounded tip, margin curved under near the base 
of the leaf, base mostly obtuse and tardily abrupt-acuminate, sometimes acute, 
long decurrent onto the petiole; lower leaf surface sparsely appressed puberu- 
lent, puncticulate, sometimes clavate glandular hairy near base, dull, brown to 
brown-green; upper leaf surface glabrous, very rarely glaucous-scintillant, pap- 
illose, puncticulate, brown-green. Venation brochidodromous; midrib promi- 
nent below, glabrous to sparsely appressed puberulent or hirtellous, epidermis 
green to brown-green, shallowly concave above, glabrous to golden hirtellous, 
sometimes partly shrouded by the lamina, then canaliculate and hairs criss- 
crossing, epidermis of basal half often black, otherwise greenish; 2° venation 
fine below, usually prominent, lateral veins 9-12 pairs, forming definite 2° 
arches with superadjacent lateral veins, arch apices 3 or more mm from the 
margin, venation tending to be obscure above, but variable; 3°-4° venation re- 
ticulated below, fine, usually apparent, but varying from obscure to prominent. 
Laminar extrafloral nectaries often up to 30, sometimes more, scattered on the 
abaxial side of the lamina, minute, peripherally rimmed with a narrow band 


292 BRIT.ORG/SIDA 22(1) 


Fic. 4, A-E. Di taricensis M.C. Provance & A.C. Sanders, sp. nov. A. Adaxial view of staminate male calyx with 
corolla removed. 8 ean pian G elypicl stamen. ie Staminiats flaws: Es aaa with leaves. A, C, and D. Based on 
Zobeida Fuentes 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 293 


of raised tissue. Male inflorescence cymes, 2-3-flowered, 1-2 cymes per axil, from 
growth of the previous year, or from the base of new growth, densely subulate 
puberulent; peduncles up to 4 mm long, sparsely to densely pubescent to stri- 
gose with light brown to black hairs, epidermis dark brown; pedicels 0.5-3.5 
mim long, vestiture and epidermis as in peduncles, pedicel bracts rarely persist- 
ing in herbarium material, 0.3-1 mm long, deltoid, densely hairy, the hairs dark- 
brown and straight. Male flowers 5-merous, calyx infundibuliform, exterior 
short-appressed gold to white puberulent, sometimes with clavate glandular 
hairs, epidermis dark brown; calyx tube 0.8-1.3 mm long, interior with ring of 
straight appressed white hairs; sepals usually erect, rarely slightly recurved, 2- 
3.5mm long, 0.8-1.5 mm wide, interior glabrous to glandular setulose, exterior 
pubescent, often densely so at the apex, the hairs wavy and amber, appearing 
glandular; corolla long-urceolate; corolla tube 3.5-7.5 mm long, 3 mm wide, in- 
terior lower-third of tube short golden pubescent, hairs + erratically directed, 
exterior evenly and densely short appressed to ascending golden hairy, also clav- 
ate glandular hairy; corolla lobes 5, quadrate to oval, 2-2.8 mm long, 1.2-2.5mm 
wide; apex sometimes bearing a long flagellate trichome, interior right mar- 
gins involute, interior glabrous, exterior appressed to ascending white puberu- 
lent, the left side of lobe densely appressed fine puberulent, sometimes with 
several minute black hairs near the apex. Stamens 17-19, adnate at various lev- 
els at and below the basal third of the corolla; filaments 1-1.5 mm long, nar- 
rowed distally, glabrous to densely erect hispidulous; anthers basifixed, 2.3-2.5 
mi long, several minute hairs at the apex, surface light-yellow apically, open- 
ing by short lateral slits confined to the distal half to two-thirds of the anther, 
sometimes opening by a complete lateral slit, pistillode + sub-conical, slightly 
wavy peripherally, + 8 long straight hairs originating from the central apex, 
otherwise glabrous. Female inflorescence flowers solitary in leaf scar axils of 
2nd year stems, or cauline at junction of previous and current growth. Fruiting 
pedicels stout, 3-18 mm long, sparsely hirtellous, reddish-gold, and gray, 
lenticellate. Fruiting calyx 5-lobed, sinuses acute; fruiting calyx tube explanate, 
sparsely appressed puberulent, scintillant, + 3.5 mm long; fruiting sepals 
accrescent, coriaceous, lorate to lanceolate, (16-)22 mm long, (5-)6-8(-9) mm 
wide, spreading to moderately reflexed, apex straight to quite incurved, 
puncticulate, shiny, golden brown, sparingly scintillant basally. Fruit berry, 
sometimes pendulant, depressed-globose, 2.5 cm tall, 4 cm wide, reportedly 
green in living material, dull, atropurpureus to dark-brown in herbarium ma- 
terial, locules 10; mesocarp reportedly yellow in living immature fruits, dark 
orange in herbarium material; hypedermis + 0.5 mm thick, stony, not wrin- 
kling in herbarium material; epidermis bullate, glaucous to pruinose-scintillant, 
either locally or over most of the fruit. Seeds not seen. 


ParatypEs: Costa Rica. Guanacaste. Canton de Liberia: Parque Nacional de G caste, Estacion Ca- 
cao, 10° 55'45"N, 85° 28'15"W, 1100 m, 24 Nov 1990, R. Espinoza 54 (kK, MO); Cordillera de G t 


294 BRIT.ORG/SIDA 22(1) 


Estacion Cacao, cna: a Estacion uMeniteas 10°55’ N, 85° 28'10" Wy, ie m, oe 996, José Gonzdlez 


et al. LIO8(K). Canton de La C 5 . V. Orosi, 
Sector Orosi oe Maritza), sendero ton Fram. 10° 57' 40"N, ne 29 45'W, 600 m, 13 Jul 1994, José 
Gonzdlez et al. 301 (MO [2 accessions]). Puntarenas. Canton de Puntarenas: Reserva Bioldgica, 


Monteverde, se de Tilaran, Altos de San Luis, Los Leitones, 10° 17'25"N, 84° 48' LO"W, 1200 m, 
4 Sep 1991, E. Bello 4014 (MO); sendero a la Catarata, por el rio, sendero Miguel Leitén, 10° 16'20"N, 84° 
49' 30"W, 1100 m, Ll Mar 1993, Zobeida ade 248(K. MO). 


Distribution and Ecology.—As far as known, this tree is endemic to northwest- 
ern Costa Rica, where it occurs in the Cordillera de Tilaran and the Cordillera 
de Guanacaste, between 600 and 1200 m in elevation. Details concerning asso- 
ciated vegetation are mostly lacking on the collections from the Cordillera de 
Guanacaste, a drier and more seasonal mountain range than the Cordillera de 
Tilaran (Hammel et al. 2004). This species could be associated with deciduous 
forests of the region. The collections from Puntarenas come from the Pacific 
side of the Cordillera de Tilaran. Label data from one of the collections, Bello 
4014, indicates that the tree was growing in charral, or young secondary forest 
(Kleinn et al. 2002). The other collection does not provide details about the as- 
sociated vegetation. Ina broad sense, the region of the occurrences in Puntarenas 
has been mapped as Costa Rican seasonal moist forests. These are deciduous 
forests that obtain 90% of their annual precipitation, (+ 1500 mm total), dur- 
ing the months of April through October (World Wildlife Fund 200D. The for- 
mation of new leaves and male flowers apparently occur during March (only 
one flowering collection was examine 

Ethnobotany.—Known by the common name ‘guacalillo’ in Puntarenas, 
Costa Rica (Fuentes 248). 

Etymology.—The epithet refers to the only country in which the species is 
currently known to occur. 

This species is different from D. conzattii in several respects. First, it is a 
buttress-forming tree attaining 35 m in height; this is quite exceptional among 
Diospyros from Central America and Mexico. Dine. conzattii is not known 
to reach over 10 m in height. The abaxial leaf venation is conspicuously brochi- 
dodromous, the lateral veins forming well defined secondary arches with the 
superadjacent lateral veins. Another interesting difference involves the distance 
from the outer-perimeter of secondary arches to the leaf margin, which is usu- 
ally around 3-6 mm at about mid-leaf. This distance is 1-3 mm in D. conzattii. 
The marginal loops of D. conzattii often become difficult, but not impossible, 
to discern, hence the aforementioned distance is measurable. Some other veg- 
etative differences useful for separating these species include yellowish to or- 
ange hairs on the adaxial midrib, and the converging adaxial lamina along the 
depressed midrib in D. costaricensis. The inflorescences of D. conzattii and D. 
costaricensis are clearly cymes, with peduncles around 6 mm and 4 mm long 
respectively. 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 295 


5. Diospyros tuxtlensis M.C. Provance & A.C. Sanders, sp. nov. (Fig. 5). Tyre: 
MEXICO. VERACRUZ: Mpio. San Andrés Tuxtla, borde de la cima del Cerro Mastagaga, al N del 
Ejido Ruiz Cortinez, Sierra de los Tuxlas, 30 May 1985, J. Calzada 11855 (HOLOTYPE: IEB-48870; 
ISOTYPE: XAL). 


Arbor usque ad 10 m alta, D. rigjae Gomez- pempe emis sed dintert tole eliptcls et non -glaucis 


marginibus non crassis et inter 
fructiferis 26-40 mm longis 


oO faa Q 
Glia ercau ans : a : 1 ea eae ay 


i 14 mm ioaeis 


Trees 8-10 m tall, probably facultatively A caenois aged terete to subterete, 
+ glabrous, half-netted to squamose, dark-brown to grey-brown with beige, 
short-fissured stem wood yellowish to orange-brown; 2nd-3rd year stems an- 
gular to subterete, glabrous, sulcate to shallow-fissured or half-netted, in com- 
binations of dark-brown, gray, and orange-brown; current year’s stems angular, 
smooth to finely sulcate, very cream to grey, shiny, lenticellate, glabrous to 
densely minute hirtellous, appressed off-white puberulent near apex, 
puncticulate. Petioles 4-7 mm long, glabrous, flat to slightly concave above, 
golden-brown, below widely rounded, glaucous-scintillant, dark-purple. Leaves 
alternate, simple, entire, pergamentaceous to chartaceous, + elliptic, 8-15 cm 
long, 3-5(-5.8) cm wide, widest at or just above the middle, both sides very 
sparsely scintillant, but leaf surface dull golden-brown to bronze between in- 
dividual scintillae, apex usually acuminate, the acumen long, narrow, obtusely 
rounded at the tip, margin flat with slight intramarginal thickening, base acumi- 
nate to attenuate, decurrent onto petiole; lower leaf surface sometimes spar- 
ingly appressed puberulent near base and midrib, conspicuously puncticulate; 
upper leaf surface glabrous, papillose. Venation arcolanguid to 
brochidodromous; midrib conspicuously flat and wide below, sulcate, purplish 
basally, becoming orange-red to golden-brown and somewhat keeled apically, 
sometimes sparsely appressed puberulent, glaucous, shallowly concave above, 
glabrous, not darkened; 2° venation raised above, obscure, lateral veins 8-9 pairs, 
shiny, chartreuse to golden-brown and conspicuous; 3° venation obscure be- 
low, granular-papillose, apparent above. Laminar extrafloral nectaries abaxial, 
common, sometimes along 2° veins, rimmed with the same bright color seen in 
2° veins Male inflorescence unknown. Female inflorescence flowers solitary, 
emerging at the junction of old and new growth. Fruiting pedicels stout, 14 mm 
long, 3mm wide, wavy-rugose, minutely fissured, lenticellate, black and tan 
mottled. Fruiting calyx with lobes and distal portion of tube strongly reflexed, 
sinuses rounded: fruiting calyx tube exterior sparsely appressed puberulent; 
fruiting sepals 5, very accrescent, 26-40 mm long, 7.5-11 mm wide, ovate to 
widely lanceolate, narrowed basally, apex acutely pointed, sometimes rounded 
sepals glabrous, golden-brown, sepal nerves distinct. Fruit berry, depressed-glo- 
bose, up to 4 cm tall, 4.5 cm in wide, atropurpureous, number of locules indis- 
cernible; mesocarp dark-brown in live material and in immature fruits of her- 
barium material; hypodermis + 0.5 mm thick, sclereidic; epidermis 


296 BRIT.ORG/SIDA 22 


1) 


Fic. 5. Diospy lensis M.C, Provance & A.C. Sanders sp. nov. Fruiting branch. Based on J./. Calzada 11855, 


glaucous-scintillant, wrinkled. Seeds immature, sizes indiscernible (broken 
seeds only), dark reddish brown, granulate to minutely ruminate (sensu Stearn 


2000, fig. 38). 


PARATYPES: Mexico, Veracruz. Mpio. San Andrés Tuxtla: senda para el Cerro Baxin, al N de San Andrés 
Tuxtla, Sierra de Los Tuxtlas, 6 Mar 1985, J. Calzada 11929 (IEB, XAL). Mpio. Soteapan: Ejido Santa 
Marta camino a la Ventana, 18° 22'N, 94° 54'W, 920 m, 19 Sep 1986, R. Acosta P & C. Gonzalez R. 1312 
(LSU, XAL). 


Distribution and Ecology.—Apparently endemic to the Sierra de Los Tuxtlas, 


PROVANCE AND SANDERS, NEW OFOLILI VE VIVO AMERICA 297 


Veracruz, Mexico. Only one elevation recorded, 920 m (R. Acosta P. & C. Gonzalez 
R. 1312). The associated vegetation types for the known collections have included 
primary selva mediana perennifolia, selva baja perennifolia, and deciduous for- 
est. It was considered common in selva baja perennifolia with elements of de- 
ciduous forest (Calzada 11855). 

Ethnobotany.—A common name has not been reported, but since the fruits 
are likely to be edible, the species may be well-known to local residents. The 
species may eventually be found as a conserved tree near homes or pastures. 

Etymology.—The epithet refers to Sierra de Los Tuxlas, Veracruz, Mexico, 
from where the only collections of this species have been made. 


SPECIES OF UNCERTAIN STATUS 


si pergamentacea Lundell, Contr. Univ. Michigan Herb. 7:44. 1942. Type. 
EXICO. CHIAPAS: Pico de Loro, near Escuintla, in advanced forest, 2200 m, 25 Jun 1941, Eizi 
ny 4278 (HOLOTYPE: MICH not seen; ISOTYPES: photo A!, photo CAS! (sterile), photo FI, IT 
not seen, photo LL-372460!, MO- 1213673) ae i 3). 
A redescription of this taxon did not seem appropriate, given that there are no 
new specimens. We do not think we can add much to the work of Lundell. Over- 
all, this species looks quite similar to D. conzattii, with which some authors 
have considered it conspecific’. This species is known only from the type speci- 
men, collected on Pico de Loro, near Escuintla, Chiapas. The petioles are very 
long and flexuous, reported to be up to 15 mm long by Lundell (1942). There 
also seem to be differences in the sepals, they being strongly reflexed, and hav- 
ing a shape that is suggestive of some populations of D. rosei sensu lato. In fact, 
these differences are significant enough that we are not entirely convinced that 
itis synonymous with D. conzattii. The leaves of the isotypes we examined had 
a thin coating of clear glue on much of their surface, which may obscure some 
characters. We recommend re-evaluating the taxonomic status of this taxon 
when additional material from the Chiapas-Guatemala borderland becomes 
available. For the time being, we do not recommend reduction to synonymy 
with D. conzattii. 


DISCUSSION 


Neither the holotype nor the two isotypes of D. riojae were available for a first- 
hand examination. Unfortunately, there are no paratypes for D. riojae, D. 
conzattii, or D. pergamentacea. Microfiche of an isotype and the detailed de- 
scription of D. riojae by Gomez-Pompa (1964) were valuable in completing this 
paper. The original description made use of tables and text in contrasting dif- 
ferences between D. riojae and D. conzattii. Additionally, it was supplemented 


>For example: A. Gomez-Pompa (1964); F. White in 1968 and C.Whitefoord & S. Knapp in 1996, in both cases by 
annotation of the isotypes. 


298 BRIT.ORG/SIDA 22(1) 


with an illustration prepared from the original collection (Gomez-Pompa, pers. 
comm.). The illustration is consistent with the type description, and the isotype 
at US, and in accordance with the Code, Article 9.2: Note 2 (Greuter et al. 2000), 
the illustration in the protologue represents original material. 

Because fruiting D. riojae material keys to D. conzattii in Standley’s treat- 
ment of Diospyros of Mexico (1924), Gomez-Pompa (1964) provided a table of 
leaf and fruit characters that could be used to separate these taxa. Admittedly, 
some of these characters overlap to some degree. When it is considered that 
Gomez-Pompa had one fruiting collection of D. riojae, one fruiting collection 
of D. pergamentacea, and probably only one collection of D. conzattii toexam- 
ine, it is to his credit that the characters he emphasized are often still useful for 
separating these taxa. We have provided an updated character table (Table 1) 
that should be helpful in the differentiation of the three new species described 
here. 

Redetermination of the available material resulted in roughly a 70% re- 
duction in the number of documented occurrences of D. riojae from what might 
have been reported based on an uncritical review of locations based on her- 
barium material. At the least, this constitutes a reaffirmation of the rarity of 
this species. The realization that D. conzattii isan element of the Veracruz flora 
is very interesting. As far as we can tell, these are the first reports of the species 
for Veracruz. Material from the Cordillera de Guanacaste, and the Cordillera de 
Tilaran, Costa Rica, represents a distinct new species, D. costaricensis. The dis- 
covery of a new species, Diospyros tuxtlensis, from the Sierra de Los Tuxtlas 
does not come as a shock, since this region is renowned for having numerous 
endemic species of plants and animals. The distributions of the closely allied 
taxa, D. tuxtlensis and D. costaricensis, are notable in light of some recent inter- 
est in Los Tuxtlas-Costa Rica disjunctions (Hammel 1997). In addition to occur- 
ring in native stands of vegetation, individuals of this taxon should be sought 
as conserved trees in local gardens and pastures. This taxon seems to be a rare 
endemic of the Sierra de Los Tuxtlas, and could be in need of formal protection. 

We have not seen female flowers of any of these species. The only descrip- 
tion of a female flower that we have seen in the literature is by Carranza (2000) 
and refers to material from Queretaro (D. gomeziorum). The lack of flowering 
Ebenaceae material in herbaria, as pointed out by Gomez-Pompa (1964) and 
Wallnofer (2001), and clearly demonstrated by our sample, makes searching 
for taxonomically informative vegetative characters a particularly attractive 
proposal (Gomez-Pompa 1964). Below we provide a key to the species described 
in this paper that emphasizes vegetative characters. Male flower and inflores- 
cence characters (Table 2) can be used to supplement the following key if de- 
sired. It should be noted that D. tuxtlensis is not treated in Table 2, since flower- 
ing material is not currently known. 


Taste 1.Vegetative and fruiting morphology across the described taxa. 


gomeziorum 


costaricensis 


conzattii 


riojae 


tuxtlensis 


Life form and size 

Leaf shape 

Leaf margin 

Leaf luster (both sides) 

Leaf vestiture below 

Leaf bloom above 

Epidermal cells of 
lamina above 

Epidermal cells of 
lamina below 


Midrib vestiture above 


Midrib topography 


above 
Number of major lateral 


veins 
Emergences on venation 


above 
Petiole color above 


Trees or shrubs to 20 m 
tall 

Feige to ovate to 
elliptic 

beveled, sometimes 


ciliolate, not thickened 


shiny 
glabrous to sparsely 
subappressed hairy 


glaucous to pruinose- 
scintillant 

not large and not 
thickene 
puncticulate 


sparsely to densely 
hirtellous to deltoid 
caly 

narrowly caniculate 


(8- 


Ww 


11-13(-14) 


2°-3° granular 
papillose 
yellow-green to olive- 
green 


Buttressed trees to 35m _ Trees or shrubs, 8-10 m 


tall 
ovate to elliptic, rarely 
widely-obovate 


curved under near base, 


not thickene 

dull 

sparsely appressed 
hairy, sometimes 
glandular 

very rarely glaucous- 
scintillant 

not large and not 
thickened 
puncticulate 


glabrous to golden 
hirtellous 


shallowly concave 
9-12 
none 


nearly black 


tall 
lanceolate to elliptic to 
ovate 


urved under near base, 
intramarginal thickening 


dull 

usually glabrous, very 

rarely glandular 

glaucous 

not large and not 
ickened 


puncticulate 


very sparsely hirtellous 


caniculate, sometimes 
raised 
9-14 


none 


pale green to green- 
brown 


Trees, rarely shrubs, to 
25m tall 
elliptic, oblong, 
obovate, or nearly oval 
cs urved, 
thickened, so 
ie 
shiny 


metimes 


glabrous 


sometimes slightly 
pruinose or scintillant 
large and thickened 


puncticulate 


glabrous, sometimes 


sparingly glandular 


hall | 
DHIAHOVWIY COMCAVE 
8-12 


2°-3° granular 
papillose 

light green to 
chartreuse to amber 


Trees, 8-10 m tall 
elliptic 


flat, slight intramarginal 
thickening 


dull 

glabrous to sparingly 
appressed puberulent 
slightly scintillant 
somewhat large and 
thickened 
conspicuously 
puncticulate 
glabrous 

shallowly concave 
8-9 

3° granular papillose 


golden-brown 


ICNIN 4N SaNnadS MAN “SHIGNYS aNv JNVAOUd 


VOIUIWY 


662 


00€ 


Taste 1. continued 


gomeziorum 


costaricensis 


conzattii 


riojae 


tuxtlensis 


Petiole color below 
Fruiting calyx posture 
Fruiting sepal apices 


Fruiting sepal length 
Fruiting sepal shape 


Fruiting pedicels 


Fruit hypodermis 


olive-green to dark 
brown, gli tening 


spreading 


apices ascending 


19-20 mm long 
lanceolate to narrowly 
elliptic or obovate 
4)5-8(-11) mm long, 
not stout 

thin 


light olive-green to very 


dark brown 


spreading to moderately 
reflexed 


apices straight to quite 


incurve 
(16-)22 mm long 
lorate to lanceolate 


3-18 mm long, stout 


thick 


pale green to green- 
brown 

spreading to slightly 
reflexed 

apices straight 


24-31 mm long 
narrow to linear 
triangular 
(7-)12-15 mm long, 
stout 

intermediate 


light green to 
chartreuse to amber 
arcuate-reflexed to 
spreadin 


oblong, narrowly 
deltoid or ovate 
7-12 mm long, stout 


thick 


dark purple 
strongly reflexed 
apices straight 
26-40 mm long 
ovate to widely 
lanceolate 


14mm long, stout 


thick 


(L)@Z VaIS/DYO"LINS 


Taste 2, Reproductive morphology of four similar taxa from Mexico and Costa Rica. 


Diospyros gomeziorum 


Diospyros costaricensis 


Diospyros conzattii 


Diospyros riojae 


Male 
inflorescence 


Male corolla 
tube shape 
Male corolla 
tube interior 
vestiture 
Stamens 


Anthers 


Filaments 


fascicles on previous year’s 
growth, solitary on new growth 


urceolate-campanulate 


sparingly hirtellous, 
hairs sometime erratic 


20 

2mm long, mer ovate 
to ovate, minutely 
granular, qe slits near 
apex 

0.3-0.8 mm long, 
ascending-puberulent 


cymes from previous year’s 
growth, cymes from new growth 


long-urceolate 


lower-third of tube 
short golden pubescent, 
hair direction erratic 
17-19 

2.3-2.5 mm long, 
several minute hairs at 
apex, short lateral slits 
in distal 2/3 to 1/2 
1-1.5 mm long, 
glabrous to densely 
erect hispidulous 


cymes from previous year’s 
growth, cymes from new growth 


long-urceolate 


sparsely puberulent, 
densest from mid-tube 
to tube bottom 

30 

3-3.5 mm long, 
lanceolate, apiculate, 
short lateral slits near 
apex 

3 mm long, minutely 
hairy 


fascicles from previous year’s 
growth, fascicles from new 
growth 


urceolate-campanulate 
deltoid scaly at mid-tube 


18 


+ 2.5 mm long, lance-ovate, 
constricted near the apex 


1 mm long, hirtellous, especially 


along margin 


ICHIN 40 CAINIAC AAIN SHIGNYS any JINVAOYd 


VoIdIWV 


LOE 


302 BRIT.ORG/SIDA 22(1) 


A KEY TO DIOSPYROS RIOJAE, DIOSPYROS CONZATTII 
AND SOME ALLIED BLACK ZAPOTES 
1. Leaves dull, laxly revolute (loosely rolled under) near the base of the leaf; neither 
the 2° nor the 3° upper leaf surface venation granular papillose. 
2. Buttressed trees up to 35 m tall and 1.5 min diameter; petiole color above et 


ae leaf margin or intramargi ina zone not thickened (viewing bottom sur- 
e); midrib ab shallowly, and often ples concave, or crimped eae 
by the adjacent sides of the lamina (boxed i gl to golden hirtel- 
lous; currently known only from Costa Rica 4. Diospyros costaricensis 


2. Trees or shrubs, 8-10 m tall, lacking buttresses as far as known; petiole color 
above pale green to green-brown; leaf margin or intramarginal zone thickened 
(on bottom surface); midrib above ately caniculate, the canal(s) raised or + 


even with the lamina; midrib very sparsely hirtellous; currently known only from 
Veracruz, Oaxaca, and possibly from Chiapas 3. Diospyros conzattii 
. Leaves shiny or dull, margins flat or sometimes subrevolute, but never loosely rolled 
under near the base; 2° or 3° venation, or both, usually granular papillose on upper 
leaf surface 
3. Leaf margin beveled outward from the upper leaf surface, down to the lower 
leaf surface;epidermal cells of the upper lamina surface not large and conspicu- 
ous with thickened anticlinal cell walls; leaves often copiously glaucous-prui- 
nose; fruiting calyx spreading, fruiting sepals with ascending sees fruiting 
edicels generally slender 2. Diospyros gomeziorum 
3. Leaf margin never beveled; epidermal cells of the upper lamina yes large 
and conspicuous, having thickened anticlinal cell walls; leaves sometimes some- 
what pruinose or scintillant, but never with copious bloom; fruiting calyx spread- 
ing to strongly reflexed, apices various; fruiting pedicels very stout 
. Leaves shiny, often tending towards oblong, oval or obovate, but may also be 
elliptic; leaf margins thickened, rounded from the top leaf surface to the bot- 
m; petiole color below light green to chartreuse or amber; fruiting sepals 
7-25 mm long, spreading to arcuate-reflexed, the apices incurved to ascend- 
ing; fruiting pedicels 7-12 mm long 1. Diospyros riojae 
. Leaves dull, elliptic; leaf margins flat or with a slight intramarginal thickening 
visible on the bottom surface; petiole dark purple below; fruiting sepals 26— 
40 mm long, strongly reflexed, the apices straight; fruiting pale 14m 
long eae tuxtlensis 


tS 


ACKNOWLEDGMENTS 


The authors are grateful to Lia DeMarco for preparing the Latin diagnoses, Clau- 
dia Ross-Ibarra for translation of the abstract in to Spanish, Edward Plummer 
for technical support and editorial assistance, Victor Steinmann for reviewing 
an early version of the manuscript, and Araceli Aguilar for her assistance. Two 
anonymous reviewers made excellent suggestions that ultimately improved the 
final manuscript. Access to microscopes was kindly made possible by Martha 
L. Orozco-Cardenas at the UCR Plant Transformation Research Center. We ex- 
tend our gratitude to the herbaria that provided material for use in this study: 
ARIZ, CAS, CHAPA, CIIDIR, DES, HUAA, IEB, K, LSU, MO, SD, UCSB, US, UTEP, 
WIS, and XAL, and also Arturo Gomez-Pompa, the Field Museum, Chicago, and 


PROVANCE AND SANDERS, NEW SPECIES OF DIOSF AMERICA 303 


the Plant Resource Center, Austin, for providing images useful in this study. 
The first author would like to extend a warm thanks to Juanita and Seychelle 
Provance for their financial support, logistic tolerance, and patience. We would 
like to thank Giles Waines and Arturo Gomez-Pompa for their insight and en- 
couragement. We are grateful to the Remib Database and Jose L. Panero (TEX, 
LL), the caretaker of the node that helped pinpoint El Cuizalin, Veracruz. Fi- 
nally, we thank UCMEXUS and the UCR Center for Conservation Biology for 
generously supporting a visit to XAL to photograph specimens. 


REFERENCES 


ANONYMOUS. 1927.Book review:Las regionas botanico-geogréficas del Estado de Oaxaca, 
Sep 1st:27-29. 

Carranza G,, E.2000. Ebenaceae. In: Fl. Bajio. Rzedowski, J.and G.Calderén R.(eds.). Instituto 
de Ecologia A.C., Centro Regional del Bajfo, Patzcuaro, Michoacan, México fasc. 83:1—9. 

Conoe, B.N. 2000. Germinacion de Diospyros riojae Gomez Pompa. Thesis. Universidad Ve- 
racruzana. México 


Gomez-Pomea, A. 1964. A new Diospyros from the Misantla region in México. J. Arnold Ar- 
bor.45:464-470. 

GreuTer, W., J. MCNeiLt, F.R. Barrie, H.M. Burbet, V. Demoutin, T.S. Fitcueiras, D.H. NicoLson, P.C. SiLvA, 
J.E. Skoc, P. TREHANE, NJ. TURLAND, and D.L. Hawkswortu (eds.). 2000. International code of 
botanical nomenclature. Regnum Veg. 138. 

Hammet, B.E. 1997. Three new species of Celastraceae from Costa Rica, one disjunct from 
Mexico. Novon 7:147-155. 

Hammel, B.E., M.H.Grayum, C. Herrera, N. Zamora (ed5.). 2004. Manual de plantas de Costa Rica: 
Volumen 1:Introduccién. Monogr. Syst. Bot. Missouri Bot. Gard. Vol. 97. Missouri Botani- 


cal Garden, St. Louis. 
Jackson, B.D. 1916. A glossary of botanic terms: with their derivation and accent. Second 
ed. London. Pp. 209. 
Keity, L.M.2001.Ebenaceae. In: Fl. Valle Tehuacan—Cuicatlan. Kelly, L.M., Ochoterena, H.,and R. 
Medina L. fasc. 34:1—5. Instituto de Biologia, Universidad Nacional Aut6noma de México. 
KeINN, C., L. Corraces, and D. Morates. 2002. Forest Area in Costa Rica: A comparative study 
of tropical forest cover estimates over time. Environ. Monit. Assessm. 73(1):1 7-40. 
Lunbett, C.L. 1942. Studies of American spermatophytes—ll. Plants of México, British Hon- 
duras, Guatemala, Costa Rica, Panama, and Puerto Rico. Contr. Univ. Michigan Herb. 


Pacueco, L. 1981. Ebenaceae. In: Fl. Veracruz. Gomez-Pompa, A. and V. Sosa, eds. fasc.16:8- 
16. Instituto Nacional de Investigaciones sobre Recourses Bidticos, Xalapa. 

Provance, M.C. and A.C. Sanpers. 2005. Diospyros torresii (Ebenaceae): a new black zapote 
from tropical Mexico. Sida 21:2045-2050. 

SANCHEZ-COELLO, N.G. 2002. Induccién de respuestas morfogenéticas en Diospyros riojae 
Gomez-Pompa en la poblacién de Cruz Blanca, mpio. de Alto Lucero, Veracruz. Thesis. 
Insituto de Genetica Forestal, Universidad Veracruzana. Xalapa, Veracruz, México 


304 BRIT.ORG/SIDA 22(1) 


Sosa, V., A.P. Vovioes, and G. Castitto-Campos. 1998, Monitoring endemic plant extinction in 
Veracruz, México. Biodiversity & Conserv. 7:1521-1527, 

STANDLEY, PC. 1922. Diospyros conzattii,a new species of persimmon from México.J.Wash. 
Acad. Sci. 12(17): 399-400. 

STANDLEY, PC. 1924. Trees and shrubs of México. Contr. U.S. Natl. Herb. 23(4):849-1312. 

STEARN, W.T. 2000. Botanical Latin, 4th ed. Portland, Oregon. Timber Press. P. 438. 

Toozia, C.A.and R.C. Keatinc. 1991. Leaf architecture of the Chloranthaceae. Ann. Missouri 
Bot. Gard. 78:476-496. 

TrasuT, D.L. 1926. Les Diospyros comestibles. Rev. Int. Bot. Appl. Agric. Trop. 6(63):675-678. 

Wattnorer, B. 2001. The biology and systematics of Ebenaceae: a review. Ann. Naturhist. 
Mus. Wien, Ser. B, Bot. Zool. 103:485-512. 

Wuite, F. 1983. Ebenaceae. In: Fl. Zambesiaca. E. Launert, ed. Vol. 7(1):248-300. 

Wauiteroorp, C. and S. Knape. 2001. Ebenaceae. In: Fl. Mesoamericana—lInternet Version, 
Davidse, G., Sousa S.,M., and S. Knapp, eds. 1998. Universidad Nacional Autonoma de 
México, México, Missouri Botanical Garden, St.Louis and The Natural History Museum, 


London. http://www. mobot.org/mobot/fm/welcome.html. 
Worto Witouire Funp. 2001. Powell, G., Palminteri, S.,and J. Schipper, preps. Costa Rican sea- 
sonal moist forests (NTO119), unrevised document. http://www.worldwildlife.org/ 
wildworld/profiles/terrestrial/nt/nt0119_full.Atml (29 June 2005 


a 


TAXONOMY OF NORTH AMERICAN SPECIES OF 
OLDENLANDIA (RUBIACEAE) 


Edward E. Terrell! Harold Robinson 
Research Associate Curator 
Department of Systematic Biology-Botany Department of Systematic Biology-Botany 
Nati ee tae n of Natural ton oe Museum of Natural History 
titution mithsonian Institution 
ee DC. 20013-7012, USA. maa DC 20013-7012, U.S.A. 
ABSTRACT 


This paper presents a taxonomic treatment of nine peas of Oldenlandia occurring as native or 
adventive in United States or Mexico. The subg Oldenlandia is represented by four species: O. 
corymbosa, O. lancifolia, O. uniflora, and O eer Subgenera for the remaining species are undeter- 
mined. Four species are native to Mexico: O. pringlei, O. microtheca, O. ovata, and O. drymarioides. A 


ninth species, O. salzmannii, is native to South domed and aavenEcy in northwest Florida and 
adjacent Alabama. Keys, descriptions, and g g f the seeds are provided. 
RESUMEN 


Este articulo presenta un tratamiento taxonomico ode nueve especies, de oo wie que ocurren 
como nativas o adventicias en Estados 
por cuatro especies: O. corymbosa, O. lancifolia, O. eer eae Los subgéneros ae la especie 
restante son eieceuen Cuatro especies son nativas de Meco O. Parnes O. microtheca, O. 
ovata y O.drymarioides. ,O. salzmannii ti 


UVOCTILICIA 
el noroeste ond y pee ae. Se proporcionan claves, pea : eee a 
arrido de las semillas. 


Oldenlandia L. (Hedyotideae; Rubiaceae) is a genus of about 100 species 
(Verdcourt 1976) distributed worldwide in warm, subtropic, and tropic regions. 
It was named by Linnaeus for Henrik Bernard Oldenlan, a Danish physician 
and botanist of the 17th century. 

Bremekamp’s monograph (1952) of the African species of Oldenlandia rec- 
ognized 61 species in 16 subgenera. Verdcourt (1976) treated 37 species of 
Oldenlandia in the Flora of Tropical East Africa, and 25 species (1989) in the 
Flora Zambesiaca region. Other authors have recognized 12 species in West 
Tropical Africa (Hepper & Keay 1963) and 7 in Gabon (Hallé 1966). Lewis (1965) 
commented that Africa has more Oldenlandia species than exist in all other 
tropical regions combined and is the center of its morphological diversity. 

enlandia was considered as only a subgenus of Hedyotis by Fosberg 
(1943) in his study of the Polynesian Hedyotis species; however, the foreign flo- 


'Address for correspondence: 14001 Wildwood Drive, Silver Spring, Maryland 20905, U.S.A 
SIDA 22(1): 305 — 329. 2006 


306 BRIT.ORG/SIDA 22(1) 


ras cited here and recent American regional floras have all treated Oldenlandia 
as a distinct genus. 

Terrell (1996) discussed problems of generic limits among Hedyotis, 
Houstonia, and Oldenlandia while revising Houstonia, a genus of 20 North 
American species. Terrell and Robinson (2003) circumscribed Hedyotis subge- 
nus Hedyotis,an Asian subgenus, and recognized the genus Exallage as a new 
subgenus of Oldenlandia. A synopsis of the United States species of 
Oldenlandia (Terrell 1990) treated five species. 

Molecular studies found Oldenlandia to be paraphyletic (Bremer 1996) or 
polyphyletic (Andersson & Rova 1999). Its species were widely distributed in 
the rubiaceous lineage in what Bremer (1996) and Bremer and Manen (2000) 
treated as the tribe Spermacoceae, that name having priority over the 
Hedyotideae within which it was nested. Tested members of Oldenlandia are 
dispersed in the slightly paraphyletic part of that lineage that Terrell and 
Wunderlin (2002), Andersson and Rova (1999) and the present authors would 
continue to treat as a separate tribe Hedyotideae. 

Lewis (1965) described the pollen of Oldenlandia and other genera in de- 
tail. In Oldenlandia the pollen are small or medium-sized and (2-)3(4-5) 
aperturate. 


MATERIALS AND METHODS 


The taxonomic treatment is based on loans of many herbarium specimens, as 
cited in the representative collections. All species except O. pringlei and O. 
drymarioides were seen in living condition. Oldenlandia microtheca and O. 
ovata were studied and collected in Mexico. Oldenlandia salzmannii was ob- 
served in greenhouse plants. The North American species of subg. Oldenlandia 
were collected in various parts of their ranges. 

Previous work on Hedyotideae has shown that capsule and seed morphol- 
ogy are especially important in taxonomic studies. In the present work we ex- 
amined the seeds of each species by scanning electron microscopy. The results 
of this study are presented below in the summary of the more significant char- 
acters of each species. 


RESULTS AND DISCUSSION 


Oldenlandia subgenus Oldenlandia 

The type species of Oldenlandia, O. corymbosa, isa worldwide weed well estab- 
lished in eastern North America. Other members of subgenus Oldenlandia in- 
clude O. lancifolia, adventive in Mexico, and the native US. species, O. uniflora 
and O. boscii. The latter two species differ from the other two in having axillary 
and terminal glomerules and slightly smaller corolla and other flower parts, 
but otherwise have similar characteristics (Table 1). 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 307 


Taste 1.Characters of species in subg. Oldenlandia. 


corymbosa lancifolia uniflora boscli 

Duration annual perennial/ann. annual perennial 
Habit erect/ prostr. erect/decumb. erect/prostr. sprdg./prostr. 
Leaves, Width mm ~9 - 2-11 1-5 
Stipules, L, mm to2 to 1.5 to4 to2 
Inflorescence pedicellate cymes pedicellate cymes glomerules glomerules 
Corollas,L, mm 1-2 15-2. 0.7-1.3 0.7-1.3 

ape rotate/tubular rotate/tubular rotate rotate 
Tube L,mm 0.5-1.0 1-0.3 0.1-0.3 
Lobes, L, mm 0.5-1.0 0.5-1.5 0.7-1.2 to 0.7 
Anthers, L, mm 0.2-0.3 0.4--0.6 0.1-0.3 0.1-0.3 
Filament, L, mm 0.1-0.3 0.2-0.5 0.2-0.4 0. 
Stig. Lobes, L, mm 0.3-1.0 0.5-1 .2 less than 0.6 less than 0.5 
Capsules, L, mm 1-2.2 1.6-3.5 1-2.5 1.5-3.0 
Seeds, L, mm 0.2-0.4 0.25-0.35 0.2-0.3 0.1-0.3 


The following description of subgenus Oldenlandia is based mainly on the 
North American species. 

Small annual or perennial herbs usually less than 80 cm tall. Stems slen- 
der, erect, decumbent, or prostrate. Leaves 3-60 mm long, opposite, sessile or 
petiolate, usually linear to ovate. Stipules 1.5-4 mm long, interpetiolar, adnate 
to leaf bases, with 1-few marginal teeth, setae, or fimbriae. Inflorescences are of 
two types, either open, pedicellate, with few-flowered cymes from axillary and 
terminal nodes or else the flowers sessile or subsessile in axillary and terminal 
glomerules. Flowers tetramerous, isostylous (homostylous) in our species, but 
species in this subgenus elsewhere may be heterostylous. Hypanthium (calyx 
cup) cup-shaped or hemispheric. Corollas 0.7-2.5(-4) mm long, rotate or tubu- 
lar, white or tinted pink or purple, glabrous externally; tube 0.1-1.0 mm long, 
shorter or longer than the 4 lobes; lobes 0.5-1.5 mm long. Anthers 0.1-0.6 mm 
long, dorsifixed, sessile or on short filaments inserted on corolla tube at or be- 
low corolla sinuses. Stigmas 2-lobed, 0.2-L2 mm long, styles filiform or thick- 
ened, glabrous. Ovules on peltate placentae. Capsules 1-3.5 x 1-4 mm, bilocu- 
lar, usually subglobose, fused with hypanthium, 3/4 to fully inferior, dehiscing 
loculicidally or also septicidally. Seeds 50-100 or more per capsule, 0.1-0.4 mm 
long, black, brown or tan, trigonous or conoidal-trigonous, basal face oval or 
elliptical, often more or less flat, lateral faces slightly or somewhat concave, 
hilum punctiform at apex of the apical angle, testa reticulate, areoles (cells) 
usually polygonal, areole walls low, rounded, indistinct, testa surface often ver- 
rucose (densely covered with minute papillae). Placenta fused with septum 
about 1/3-1/4 of the distance above its base. Chromosome number x = 9. 

When compared with the other species (Table 2) the unique or unusual 


308 


Taste 2. Comparison of four Oldenlandia taxa. 


BRIT.ORG/SIDA 22(1) 


subg. Oldenlandia Pringlei Salzmannii Microtheca 
No.of species examined four one one three 
Habit erect/prostrate ascdq./prostr. creeping erect/decumbent 
rhizomatous 
Leaves Lxw,mm 3-60 * 1-12 3-15 XK 03-2 1.5-5.2 X 0.7-3 3-40 * 1-20 
Stipules L 1.5-4mm 0.3-1.0 to 0.5 0.5-2 
Inflorescence cymose/glomer. cymose solitary cymose 
Corollas L - 4.0-8.5 2-5.5 25-7 
Cor. shape rotate/tubular subsalv./funnel.  subsalverform — funnelform 
Cor. tube L 0.1-1.0 mm 1-2.2 1-3 
Cor. lobes L 0.5-1.5 mm 1.5-3.5 1-3.2 1.5-4 
Anthers L 0.1-0.6 mm 0.8-1.3 0.5-0.8 0.4-1 
Capsules L x W 1—3.5 x 1-4 2-3 X 2-2.5 1.5 X 1.5 1-3 X 1.5-3.5 
Seeds no./caps. 50-100 + 30-50 4-14 10-34 
Seeds L/dia.mm 0.1-0.4 2-0. 0.3-0.5 0.4-0.7 
Seeds shape trigonous subglob/ovoid trigonous angulate/irreg 
conoidal 
Seeds, areoles polygonal polygonal polygonal none 
Areole walls low, rounded thick, sinuous distinct none 
Testa reticulate reticulate reticulate not reticulate 
Testa surface verrucose/other — verrucose smooth entangled 
str 
Chrom.no. x= 9 ? 15 11,12 


features of subg. Oldenlandia include the following: Inflorescence either pedicel- 
late in small cymes at the nodes or else flowers subsessile in glomerules; flowers 
usually isostylous (homostylous); corollas small, often 0.7-2.5 mm long, rotate 
or tubular; anthers 0.1-1.0 mm long; seeds 50-100 or more per capsule, 0.1-0.4 
mm long, trigonous, areole walls low, rounded, indistinct; chromosome num- 
ber x =9 in many species worldwide. 

Seed data for this subgenus are described and illustrated in Figures 1-3. 
Figure | shows typical trigonous oldenlandioid seeds in O. corymbosa and O. 
lancifolia witha reticulate surface composed of polygonal areoles. A basal face 
is often flat (Fig. 1C) and the lateral faces are flat or slightly concave. The apical 
hilum is shown in Fig. 1D. An enlargement of part of an areole (Fig. IF) has a 
verrucose testa with low areole walls. 

Two Linnaean species, the Asian O. herbacea and the African O. umbellata 
(Fig. 2) show trigonous seeds with concave lateral faces lacking the verrucose 
surface; instead with a coarsely papillose (Fig. 2C) and an apparently smooth 
surface (Fig. 2D). 

Oldenlandia uniflora has trigonous seeds (Fig. 3) with a concave lateral 
face (Fig. 3B,C), a densely papillose surface and low indistinct areole walls. For 


td 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 309 


Fic. 1. Seeds of Oldenlandia speci ined by SEM. A—D. O/deniandia corymbosa, A. Proctor 10493 (US), Jamaica; B, D. 
Lelong 6891.2 (NCU), Florida; C. Standley 52679 (US), Honduras. E-F. Olden/andia lancifolia, Vasquez et al. V-1324 (XAL), 
Mexico. A. Views of 5 seeds; B, E. End views; C. Tilted showing basal and lateral surfaces; D. Hilum at apex; F. Areoles 


enlarged. 


310 BRIT.ORG/SIDA 22(1) 


Fic. 2. Seeds of O/denlandia species examined by SEM. A, C. Oldenlandia herbacea, Fosberg 40749 (US), Nigeria. B, D. 
Oldentandia umbellata, Fosberg 51929 (US), Ceylon. A—B. End views; C—D. Areoles enlarged. 


comparison with seeds of subgenus Oldenlandia two trigonous seeds are shown 

(Fig. 3D,E) from the Asian species, O. lapeyrousii, belonging to the subgenus 

Exallage lerrell & Robinson 2003). 

Oldenlandia species not in subgenus Oldenlandia 

Table 2 summarizes the more important characters of the remaining species as 

well as those of subgenus Oldenlandia. The remaining species are not readily 

assignable to new subgenera until more data are known about other genera. 
Oldenlandia pringlei, is restricted to San Luis Potosi, Mexico. Its more sig- 

nificant characters are as follows: Plants small, herbaceous, rhizomatous, as- 

cending or prostrate; leaves linear; flowers in open, few-flowered cymes, 


heterostylous; corollas 4+.0-8.5 mm long, subsalverform in contrast with the 
small, rotate or tubular corollas present in subg. Oldenlandia; anthers 0.8-1.3 
mm long; seeds 30-50 per capsule, subglobose, ellipsoid, or ovoid instead of 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 311 


Fic. 3. Seeds of Oldenlandi ined by SEM. A—C. Oldenlandia uniflora, Leonard 7450 (FSU), South Carolina. D- 


E. Ofdenlandia lapeyrousii, Smith 9079 (US), Fiji. A, D, E, end views; B-C. Two views of enlarged areoles. 


trigonous, and areole wall rather thick and sinuous, chromosome number not 
known. Ventral and dorsal views of the ellipsoid seeds with their sinuous thick- 
walled areoles are shown in Figure 4 A,B. The ventral view shows the centric 
punctate hilum. 

Oldenlandia salzmannii is a South American species established in north- 
western Florida and adjacent Alabama. The plants are herbaceous, creeping 
perennials, whereas other Oldenlandia species may be prostrate but rarely or 
not creeping; leaves small, 1.5-5.2 mm long; stipules small, to 0.5 mm long; flow- 
ers solitary, pedicellate; corollas subsalverform; capsules small, 1.5 < 1.5 mm; 
seeds only 4-14 e, 0.3-0.5 mm long, trigonous, areole walls distinct; 


peer 
— 


per capsu 


312 BRIT.ORG/SIDA 22(1) 


Fic. 4. Seeds of Oldenlandia species examined by SEM. A-B. Oldenlandia pringlei, Pringle 3758 (US), Mexico. C-F. 
Oldenlandia salzmannii, Burkhalter & Hand 6537 (UWFB), Florida. A. ventral view; B. dorsal view; C. side view; D. hilar 
area; E. basal surface; F. areoles. 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 313 


chromosome number n = 15; 2n = 30, a unique number in Hedyotideae (Lewis 
1966b). Figure 4 C-E shows a trigonous seed with concave lateral faces, an api- 
cal hilum, and polygonal areoles with straight walls. 

The Oldenlandia microtheca group includes three Mexican species, O. 
microtheca, O. ovata, and O. drymarioides. Their relationships to other species 
or groups are somewhat in question, but they have been named Oldenlandia 
and appear closer to that genus than to any other. Oldenlandia drymarioides is 
a rarely collected species with a close resemblance to O. ovata. It is tentatively 
considered a distinct species. 

The characteristics (Table 2) of the three species may be summarized as 
follows: Small herbs, annual or perennial; flowers in cymes, heterostylous; co- 
rollas 2.5-7.0 mm long, funnelform; capsules 1-3 mm long; seeds 10-34 per cap- 
sule, 0.4-0.7 mm long, angulate or irregularly conoidal, surface irregularly hon- 
eycombed or alveolate, areoles lacking, coalesced, replaced by entangled 
vermiform strands; chromosome number x = 11, 12. These species are notable 
for their unusual seeds (Fig. 5), which differ conspicuously from those of all 
other Oldenlandia species. The seeds lack the usual reticulate testa and instead 
have a jumbled mass of entangled vermiform strands, a condition here referred 
to as coalescent areoles in reference to their running together (Fig. 5 C.F). These 
seeds are larger and fewer per capsule as opposed to subgenus Oldenlandia and 
show a range of shapes from irregularly and obtusely angulate to irregularly 
conoidal. Coalescent areoles and a chromosome number of x= 11 are known in 
Houstonia subgenus Chamisme section Ericotis of southwestern U. S. and 
Mexico (Terrell 1996), but those species are otherwise quite distinct from the O. 
microtheca group. 


_ 


TAXONOMIC TREATMENT OF NORTH AMERICAN SPECIES 


Oldenlandia L., Sp. Pl. 119.1753. Gerontogea Cham. & Schlecht. Linnaea 4:154.1829. TyPE: O. 
corymbosa L., designated by Hitchcock and Green (1929). Hedyotis ae bosa (L.) Lam., Tabl. 
Encycl. 1:272. 1792. Lectotype: Plumier, Nov. Pl. Amer. t.36. 1703, cited by Jarvis et al. (1993) 

and Verdcourt (1976). Four other generic synonyms were listed by eae (1918) 


KEY TO UNITED STATES SPECIES OF OLDENLANDIA, NATIVE OR ADVENTIVE 


1. Creeping, mat-forming perennials; leaves 1.5-5.2 mm long; adventive in northwest 

Florida and adjacent Alabama 6. O. salzmannii 

Erect, spreading, decumbent, or prostrate annuals or perennials; leaves (3—)5— 

mm long; native or widely established species. 

2. Flowers and capsules on pedicels more than 3 mm long, not in glomerules 
1.0.corymbosa 

2. Flowers and capsules in axillary or terminal glomerules, sessile or pedicels less 


= 


3. Annual:leaves 2-11 mm wide, ovate to elliptic 3.0. uniflora 
3. Perennial with woody tap root; leaves1—3(-5) mm wide, linear to narrowly 
elliptic 4.0. boscii 


314 BRIT.ORG/SIDA 22(1) 


Fic. 5. Seeds of Oldenlandia ae) Semines by oe A- 2 eae peli Mg B. Schiede 390 (K), Mexico; C. 
Pennell 17920 (US), Mexico. D—F | Mexico. A—B. side views; D-E. 


ventral views including hilar areas; C—F. pen of coalesced areoles. 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 315 


KEY TO MEXICAN SPECIES OF OLDENLANDIA, NATIVE OR ADVENTIVE 


1. Corollas 0.7-2.5 mm long, rotate or tubular; anthers 0.2-0.6 mm long;seeds 0.1—0.4 
mm long, trigonous. 
2. Plants often 10-24 cm tall; inflorescence with 1-6 pedicels per node, pedicels 
usually 2-13 mm long; anthers 0.2-0.3 mm long; capsules 1-2.2 X mm 
locules not saccate, apices truncate or retuse 1.0. corymbosa 
2. Plants often 15-60 cm tall; inflorescence with 1(-3) pedicels per node, pedicels 
(-30) mm long ; anthers 0.4-0.6 mm long; capsules 2-3.5 X 2.5-3.5(-4 
mm, locules appearing inflated or saccate, apices rounded or beaked 2.0. lancifolia 
. Corollas 2.5-8.5 mm long, subsalverform or funnelform; anthers 0.4-1.3 mm long 
seeds 0.2-0.7 mm long, subglobose, ovoid, or irregularly angulate. 
3. Plants often with rhizomes; leaves 0.3-2 mm wide; seeds subglobose, ellipsoid, 
or ovoid, testa reticulate, areoles polygonal; endemic to San Luis Potosi, Mexico 
5.0. pringlei 
3, Plants without rhizomes; leaves 1-20 mm wide; seeds angulate or irregularly 
conoidal, testa not reticulate, areoles lacking, coalescent, replaced by entangled 


— 


trands., 
4, Perennials; plants 9-42 cm tall; leaves 2-20 mm wide; flowers heterostylous 
7,0. microtheca 


4. Annuals, plants less than 20 cm tall; leaves 1-9.5 mm wide; flowers isostylous. 
5. Basal leaves absent or smaller than cauline 8.0. ovata 
5. Basal leaves larger than cauline 9.0. drymarioides 


1. Oldenlandia corymbosa |. Sp.P1.119.1753. Hedyotis corymbosa (L.) Lam, Tab. Eneycl. 

72.1792. TyPE: Plumier, Nov. PI. Amer. Gen. 42, t.36.1703. (Verdcourt 1976; Jarvis et al. 1993). 
Small annual herb. Stems 10-24(-40) cm tall, slender, tetragonal, erect, spread- 
ing, decumbent, or prostrate, glabrous or puberulent toward base or at nodes, 
with l-many branches. Leaves (5-)10-40 x 1-7(-9) mm, sessile or short-peti- 
olate, l-nerved, narrowly elliptic, elliptic, narrowly oblong, or linear, glabrous, 
margins glabrous or ciliolate near leaf bases, apices acute or apiculate. Stipules 
to 2 mm long, to 3 mm wide, whitish, truncate or rounded, with 0-few mar- 
ginal teeth or setae to 3mm long. Inflorescence cymose, flowers isostylous, pe- 
duncles absent or present, similar to pedicels, usually 5-15 mm long, filiform, 
pedicels from most nodes, 1-6 per node, usually 2-13 mm long, filiform. Hy- 
panthium glabrous or scabrous; calyx lobes 0.5-1.3 mm long, slightly exceed- 
ing capsule, lanceolate, sometimes ciliolate on margins and sinuses, acute. Co- 
rollas 1-2 mm long, rotate or tubular, white or occasionally faint lavender or 
pink, partly obscured by calyx lobes; tubes 0.5-1 mm long, throat with ring of 
white, sometimes clavate, hairs; lobes 0.5-1 mm long, ovate, spreading. Anthers 
0.2-0.3 mm long, ovate, whitish or purplish, on filaments 0.1-0.3 mm long, at- 
tached to sinuses of corolla tube. Stigma lobes 0.3-1.0 mm long, style thickened. 
Capsules 1-2.2 x 1.3-2.8 mm, subglobose or slightly wider than long, not or 
slightly compressed, glabrous, 4/5 to fully inferior, thin-walled, fragile, apices 
truncate or retuse. Seeds 50-100 or more per capsule, 0.2-0.4 mm long, usually 
brown, trigonous, hilum apical, punctiform, testa reticulate, areoles polygonal, 


316 BRIT.ORG/SIDA 22(1) 


their walls low, rounded, indistinct, walls and testa surface densely covered with 
minute papillae. Flowering all year in tropical climates. Chromosome number: 
n= 9,18, 27: 2n = 18, 36, 54 (Lewis 1964, 1965, 1966c). 

Distribution and habitats—Pantropic weed. United States: adventive and 
becoming more common in Atlantic and Gulf Coastal Plains and Mississippi 
Embayment. North Carolina, South Carolina, Georgia, throughout Florida, 
southern parts of Alabama, Mississippi, and Louisiana, to eastern Texas. Mexico: 
Nayarit, Tabasco, Chiapas (probably more frequent in Mexico than indicated 
by available records). Standley (1918) did not give any records for Mexico and 
United States, listing only West Indies, Central America, and South America in 
the Western Hemisphere. Hawaii: Oahu, Hawaii, Maui. Eastern Hemisphere. 
Disturbed places, lawns, roadsides. 

Discussion.The description refers to the diploid race of var. corymbosa. 

Further comments on varieties in Lewis 1965, 1966c, Verdcourt 1976; Sivarajan 
and Biju 1990. Specimens cited below are considered to be var. corymbosa. 
. collections. U.S.A. FLORIDA. Alachua:, lawn, McCarty Hall, ie! of Florida, 
Gainesville, D’Arcy 2160 (FLAS). Collier: | mi S of Naples, ne et al. 9071 (USF). Dade: Homestead, 
Radford & Leonar fei ). Escambia: West Brainerd St, near Pensacola, Burk aa “LAS). 
Glades: 3 mi SW of Palmdale, Ward 5185 (FLAS). Hillsborough: oF of International Airport, Tampa, 
Lakela 24642 ee USF). Manatee: Bradenton, Cuthbert 1442 (FLAS). Orange: Orlando, Schallert 
1590 (FLAS). Palm Beach: 836 Biscayne Drive, West Palm Beach, Cassen 559 (FLAS). Pinellas: Gulf- 
port, Thorne 13874 (US). Sarasota: opposite entrance to Pine Park west of US 441, Ward & Burch 3110 
(FLAS). LOUISIANA. Orleans: Tulane University campus, Sundell 2028 (NO). SOUTH CAROLINA. 
Florence: Wynn Owens a Evergreen, Swails 83/971 (USCH). Richland: 6511 Helena Road, Colum- 
bia, Nelson 402 (FLAS) 


2. Oldenlandia lancifolia (Schumach.) DC., Prodr. 4:425.1830. Hedyotis lancifolia 
Schumach. [in Schumach. & Thonn], Beskr. Guin. Pl. 72.1827. Type: GHANA: Valley of 
Aquapim, Thonning 210 (LECTOTYPE: C; ISOLECTOTYPE: S, n.v.). 

Hedyotis commutata Schult. & Schult. f., Mant.3:134.1827. TyPE: PUERTO RICO: Bertero s.n., nv. 


Perennial herb (rarely annual). Stems 15-60 cm tall, slender, sometimes flat- 
tened after drying, erect, decumbent or sprawling on other vegetation, often 
rooting at base, glabrous to densely puberulent. Leaves 10-60 x 2-12 mm, sessile 
or subsessile, l-nerved, linear, lanceolate, or elliptic, glabrous, margin revolute, 
apices acute. Stipules to 1.5 mm long, to 2.5 mm wide, whitish, truncate or 
rounded, with 1-few marginal teeth or setae to 3-4 mm long. Inflorescence cy- 
mose, flowers isostylous, pedicels from most nodes, 1-2(-5) per node, 5-15(-30) 
mm long. Hypanthium glabrous or pubescent; calyx lobes 0.7-L6 mm long, 
exceeding capsules, lanceolate or deltate, scaberulous to puberulent, acute. Co- 
rollas 1.0-2.5 mm long, rotate or tubular, white; tube 0.5-1 mm long; lobes 0.5- 
1.5 mm long, usually slightly longer than tube, ovate, spreading. Anthers 0.4- 
0.6 mm long, oblong, on filaments 0.2-0.5 mm long exserted ca. 0.5mm beyond 
throat, attached at sinuses of corolla. Stigmas lobes 0.5-1.2 mm long, linear, 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 317 


exserted ca 1 mm beyond throat, styles 0.5-1 mm long, filiform. Capsules 1.6-3.5 
x 2.5-4 mm, subglobose or slightly wider than long, slightly compressed, gla- 
brous, 3/4 to 9/10 inferior, thin-walled, fragile, apices rounded or somewhat 
beaked especially after dehiscence, locules appearing saccate. Seeds numerous 
per capsule, 0.25-0.35 mm long, trigonous, hilum apical, punctiform, testa re- 
ticulate, areoles polygonal, their walls low, rounded, indistinct, walls and testa 
surface densely covered with minute papillae. Chromosome number: n= 18, 2n 
= 36 (Lewis 1965). 

Distribution and habitat.—Tropical Africa; introduced or adventive in 
South America, Central America, West Indies, and Mexico. Recorded from 
Mexico in Tamaulipas, Veracruz, Oaxaca, Chiapas, and Tabasco. Low altitudes 
in moist places along streams, swamps, forest openings. 

Discussion.—Verdcourt (1976) provided data on African varieties of O. 
lancifolia. 

Many American collections of this species have been misidentified as O. 
herbacea (L.) Roxb, an Asian species; see key below. The occurrence of any O. 
herbacea collections in Mexico has not been substantiated by us. 


KEY TO OLDENLANDIA LANCIFOLIA AND O. HERBACEA 


1. Stems weak, often reclining on other eed sometimes flattened in drying; 


leaves 2-12 mm wide; corollas 1.0-2.5 mm long; capsules 2.5-4 mm wide, 
subglobose or wider than long, locules a capsules often appearing inflated or 
saccate O. lancifolia 
ee erect, solid, tetragonal; leaves 1-4 mm wide; corollas 2.5-4.7 mm long; cap- 
sules 1.5-2.5 mm wide, subglobose or longer than wide, locules not appearing 
ee or saccate O. herbacea 


Both Hedyotis commutata and H. lancifolia were created in 1827, and apparently 
have equal priority. The epithet lancifolia has been the choice of Verdcourt 
(1976), Howard (1989), and others in floras. 


Representative specimens. COSTA RICA. Cartago: Rio Reventazon, Catie Agric. Station, Turrialba, 
Hillet al. 17735 (HILL). San Jose: Vic. ne General, Skutch 3932 (GH, MO, US). Limon: Vic. of Guapiles, 
iid 37296 (US), GUATEMALA. Izab c. of Quirigua, penta 24267 (GH, US). HONDURAS. 
mayagua: Vic. of tae ai sane a ons US). MARTINIQUE. 1 1/2 km S of Ajoupa-Bouil- 
ns Kimber 687 (WIS). MEXICO s: Mpio. of see i Agua, W of ee on road to 
Villahermosa, Breedlove e ute 5228 (CAS, MICH, TEX). Tamaulipas: 8 mi from Tampico on 
Mante highway, Jol tham 4077 (MEXU, MICH, TEX). Veracruz: Rio Sea aorillas del 
Campamento nos Cedillo, Haalgeetlen Brigada Vasquez 172 (XAL). NICUARAGUA. Zelaya: Pu- 
erto Isabel, Narvaez S. 2890 (GH). PANAMA. Bocas del Toro, Changuinola to 5 mi S$ at jet. of Rios 
Changuinola and Terebe, Lewis et al. 796 (GH). 


3. Oldenlandia uniflora es SP. Bie 91753: aaa unilona (L.) Lam.,Tab.Encycl. 
272. Ed i ) Raf., Actes Soc. Lin 9.1834. TYPE: VIRGINIA: 
Clayton 587 (LINN -155. 3) ISOTYPE: BM!). 


Hedyotis auricularia Walter, Fl. Carol.85.1788. (non H. auricularia L., Sp. PL. 101.1753). TYPE: n.v. 
Oldenlandia glomerata Michx., Fl. Bor.-Amer. 1:83. 1803. Hedyotis glomerata (Michx.) Elliott, 


318 BRIT.ORG/SIDA 22(1) 


Sketch Bot. S. Carolina 1:188. 1816. Stelmotis glomerata (Michx.) Raf., New FI. 4:10L. 1830. 
Stelmanisglomerata (Michx.) Raf., Autik. Bot. 13.1840. Type: “in humidis Carolinae inferioris,” 


nv, 
Hedyotis virginica Spreng., Pl. Min. Cogn. Pug. 2:34. 1815. TYPE: n 
Hedyotis fasciculata Bertol., Mem.Reale Accad. Sci. Ist. Bologna 2: oi 1850.0. fasciculata (Bertol.) 
small, FIS.E.U.S. 1106. 1903. H. uniflora var fasciculata (Bertol.) WH. Lewis, Amer. J. Bot. +9:865. 
1962. TYPE: nv. 
Oldenlandia aa C. Mohr, Bull. Torrey Bot. Club 24:27. 1897. Type: ALABAMA: Mobile, low 
wet places, borders of ponds, Grand Bay [near Mobile], Sep 6, 1888, C. Mohr s.n. (LECTOTYPE, 


here designated: UN 


Small annual herb. Stems 2-70 cm tall, slender, erect, spreading, decumbent, or 
prostrate, eee white-hirsute or villous to glabrous, much branched. Leaves 
3-20(-28) x 2-11 mm, sessile or short-petiolate (to 2 mm), ovate or elliptic, hir- 
sutulous or pubescent to glabrous above or pubescent only on midribs and 
nerves, glabrous or pubescent on nerves beneath, margins ciliolate to glabrous, 
apices acute or obtuse. Stipules to + mm long, to 3mm wide, whitish, divided 
near base into l-few (sometimes branched) ciliate, linear or lanceolate, curved 
or straight teeth or setae toca 5mm long. Inflorescence with |-many isostylous 
flowers in axillary and terminal glomerules, sessile or on pedicels to ca 3mm 
long, sometimes flowering when plants are very small. Hypanthium hirsute to 
glabrous; calyx lobes 0.8-2.5(-4) x 0.5-2 mm, strongly one-nerved near base, 
ovate or ovate-lanceolate, pubescent and ciliate to glabrous. Corollas 0.8-1.3 mm 
long, rotate, white or pale blue; tube 0.1-0.3 mm long; lobes 0.7-L.2 x 05-11 
mm, usually shorter than the calyx lobes, ovate. Anthers 0.1-0.3 mm long, ovate, 
purple, on filaments 0.2-0.4 mm long attached to corolla tube at sinuses. Style 
and stigma 0.2-0.6 mim long, stigma lobes included in tube, style thickened. 
Capsules 1-2.5 x 1-3 mm, subglobose or slightly wider than long, densely hir- 
sute with whitish hairs or pubescent to glabrous, 7/8 or fully inferior, walls 
medium thick, apices truncate, dehiscence loculicidal. Seeds numerous per cap- 


sule, 0.2-0.3 mm long, black, brown, or tan, trigonous, hilum apical, punctiform, 
testa reticulate, areoles polygonal, their walls low, rounded, indistinct, walls and 
testa surface densely covered with papillae. Flowering in United States in spring 
(Florida), summer, fall. Chromosome number: n = 18, 36; 2n = 36 (Lewis 1962). 

Distribution and habitats—United States: Mainly in Atlantic and Gulf 
Coastal Plains and Mississippi Embayment. New York (Long Island) south to 
southern Florida, west to eastern Oklahoma, eastern Texas, southern Arkansas, 
southeastern Missouri, western Kentucky, and western Tennessee. West Indies 
in Cuba, Puerto Rico, and Jamaica. Not recorded from Mexico. Wet or moist 
places, lake shores, swamps, stream banks and sand bars, pine and deciduous 
woods (often in openings), savannahs, fields, roadsides, gravel pits. 

Discussion.—Variation in leaf shape and plant vesture does not appear to 
be taxonomically significant. This species produces flowers at an early age: one 
collection from Virginia was flowering when plants were 4 cm tall. 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 319 


apenas collections. CUBA. Isle of Pines: San Pedro and vicinity, Britton & Wilson 14703 (CM). 

DA. Glades: 8.6 mi SE of Palmdale, Lewis 5682 (TEX). Highlands: 6.7 mi S of US 27 and 
Florida ae intersection, Ray et al. 10420 (USF). Manatee: 2 1/2 mi N of Duette and road 62, Shuey 2113 
(DS). Sarasota: 6 mi SE of Sarasota, Ward & Burch 3196 (MO). Walton: 4 mi S of Freeport, Davis & 
Davis eee (CM). GEORGIA. McIntosh: NE part of Sapelo Island, Duncan & Adams 17858 (MICH). 
LO . Beauregard: 1 1/2 mi Sof Bayou Amacoco, Ewan 20089 (NO). East Baton Rouge: Magno- 
lia, rate ene U). St. Tammany: near jct of hwys. 1-10 and I-59, Darwin & Sundell 918 (NO). MARY- 
D. Prince Georges: Magruder Landing, Smith 5131 (MT). MISSISSIPPI. Hancock: 0.5 mi W of St. 
Louis Bay and hwy. 90, Lewis 5688 (TEX). NEW YORK. Long Island: near Long Pond, Wading River, 
Miller s.n., 8/23/1878 (CM). NORTH CAROLINA. Hoke: 4 mi SSW of Ashley Hts., Ahles 36373 (ASU, 
CM, MICH). OKLAHOMA. Atoka: 0.5 mi NW of Boehler, Taylor 20365 (KANU). SOUTH CAROLINA. 
Aiken: Montmorenci, Ahles & hae 55058 (ARIZ, ASU, BALT, CLEM, CM, DS, KANU, KY, LL, 
MICH, TEX, US, VPI). TEXAS. Cherokee: Larissa, Palmer 8613 (CAS). Nacogdoches: 3.5 mi S of 
Nacogdoches, Lewis & Oliver 5510 (Ds, TEX). VIRGINIA. Accomac: Parksley, Norton s.n. 9/11/02 
(MARY) 


oO. 


Bs 
Z 


4. Oldenlandia boscii (DC.) Chapm., Fl. South.U.S.181.1860. Hedyotis boscii DC., Prodr. 
4:420. 1830. Tyre: U.S.A: Carolina, 1798-1800, Bosc s.n. (G-DC (n.v.): microfiche US). 


Small perennial herb with woody taproot to ca 6 mm thick. Stems often nu- 
merous, 5-30 cm tall, slender, spreading, decumbent or prostrate, forming mats 
or mounds toca 4+ dm wide, glabrous to minutely papillose or puberulent, much 
branched. Leaves 10-25(-30) x 1-3(-5) mm, sessile, linear to narrowly elliptic, 
glabrous to minutely papillose or puberulent above, glabrous or scabrous on 
midrib beneath, margins glabrous or scabrous, sometimes revolute, apices ob- 
tuse or acute. Stipules to 2 mm long and wide, whitish, truncate or deltate, with 
l-several marginal teeth or setae to 2 mm long. Inflorescence with l-several (- 
many) isostylous flowers in axillary and terminal glomerules, sessile or 
subsessile. Hypanthium glabrous or minutely papillose; calyx lobes 0.8-2 x 
0.3-0.8 mm, strongly l-nerved, deltate to lanceolate. Corollas 0.8-1.3 mm long, 
rotate, white, pink, or lavender; tube 0.1-0.3 mm long; lobes 0.7-1L.2 mm long, 
ca. 1-2 mm wide, shorter than the calyx lobes, ovate. Anthers 0.1-0.3 mm long, 
broadly elliptic, on filaments ca. 0.2 mm long attached to corolla at sinuses. 
Stigma lobes included in tube, style and stigma less than 0.5 mm long, style 
thickened. Capsules 1.5-3 x 1.5-2.5mm, subglobose or often slightly longer than 
wide, verrucose or papillose varying to glabrate, truncate or retuse, 7/8 to fully 
inferior, walls medium thick, dehiscence loculicidal. Seeds numerous per cap- 
sule (to at least 120), 0.1-0.3 mm long, black or brown, trigonous, hilum apical, 
punctiform, testa reticulate, areoles polygonal, their walls low, rounded, indis- 
tinct, walls and testa surface densely papillose. Flowering April to November. 
Chromosome number: 2n = 36, based on three collections (Lewis 1962). 
Distribution and habitats —United States: Mainly in the Atlantic and Gulf 
Coastal Plains, Mississippi Embayment, and Ozarks in the southeastern states: 
Southeastern Virginia, North Carolina, South Carolina, Georgia, northern 
Florida, and west to central and western Tennessee, southeastern Missouri, east- 


320 BRIT.ORG/SIDA 22(1) 


ern Oklahoma, and eastern Texas. Wet or moist places, stream banks, lake shores, 
roadsides, ditches, fields, woods, savannahs, disturbed open places. 


Representative collections. U.S.A. ALABAMA. Escambia: Conecuh River bottoms 6 mi E of Boykin on 
US 29, Kral 44857 (K ANU). ARKANSAS. Faulkner: Conway, Haas 678 (US). FLORIDA. Gadsden: Flood- 
plain Appalachicola River at Chattahoochee, Godfrey 67526 (FLAS, TEX, USF). Leon: 10 mi S of Talla- 
gee on Spring Hill Road, Godfrey 62904 (MT, TEX, USF). Suwannee: Live Oak, Curtiss 6649 (ISC, 
US). GEORGIA. Miller: 7 mi ESE of Colquitt, Thorne 5199 (MT, US). LOUISIANA. Allen: 7.2 mi W of 
re Shinners 21522 (MICH). Grant: Louisiana road 8 ca 41/2 mi NE of Colfax, Allen et al. 7802 
(NO). Sabine: Zwalle, Toledo Bend Reservoir, Sabine River, Demaree 48352 (NO). Washington: Along 
Pearl River E of Angie, Brown 17866 (LSU). MISSISSIPPI. Jackson: Biloxi, Tracy 6422 (US). Oktibbeha: 
10 mi Sof Starkville, McDaniel 2661 (NO). NORTH CAROLINA. Scotland: Sink Hole Bay, SF side of SR 
1622, Berg 1156 (NCU). OKLAHOMA. McCurtain: | mi N of Bethel, Waterfall 14776 (DUKE). SOUTH 
CAROLINA. Aiken: Montmorenci, Ahles & Crutchfield 55054 (ASU, CM, FLAS, KANU, KY, MICH, 
TEX, USCH, VPI. TEXAS. Bastrop: Bastrop, Tharp s.n., 8/7/38 (CAS, DS, MICH, US). Jefferson: Beau- 
mont, Palmer 12720 (US). Nucces: Padre Island, Jones 2244 (WWE). VIRGINIA. Southampton: Predler’s 
Pond, Nottoway Swamp, SW of Sedley, Fernald & Long 7625 (US). Sussex: Airfield Millpond, SW of 
Wakefield, Fernald & Long i (US). 


5. oneal prnsiet BL. HOR Proc. se aca. Arts 27169. eee Hedyotis 


line eins Halicnde de nes a, 7 Jun 189], CG. eile! 3758 ( (LECTOTYPE, fae ee 

nated: GH: SOL ECTOTYPES: BMI, Fl, K!, MICH!, MO!, MSC, PHI, US-3), VT! 
Small perennial herb from slender scaly rhizomes and slender tap root. Stems 
5-20 cm tall, slender, subterete, scaly toward base, ascending or prostrate, 
densely puberulent to glabrate, branched from base or upper nodes. Leaves 3- 
15 x 0.3-2 mm, thickened, slightly fleshy, sessile, linear, glabrous or granular- 
puberulent. Stipules usually 0.3-1.0 mm long, somewhat sheathing, deltate or 
ovate, with marginal gland-tipped teeth or lobes.Inflorescence in terminal or 
axillary few-flowered cymes, pedicels to ca. 10 mm long, flowers heterostylous. 
Hypanthium glabrous; calyx lobes to 2 mm long, 0.2-1.0 mm wide, lanceolate 
or ovate-lanceolate, obtuse. Corollas 4.0-8.5 mm long, subsalverform or nar- 
rowly funnelform, white, with dark nerves; tube 2-5 mm long, 0.7-2 mm wide 
at throat, slightly widened distally, glabrous within proximally, puberulent 
distally; lobes 1.5-3.5  0.7-1.3 mm, lanceolate-ovate, puberulent within. An- 
thers 0.8-1.4 mm long, narrowly oblong. Stigma lobes 1-1.3 mm long, subglobose. 
Pin flowers with stigmas exserted to 2.5 mm beyond throat; anthers included, 
filaments 0.51.0, attached near midpoint of tube to just below throat. Thrum 
flowers with anthers sessile or on short (0.5 mm) filaments, exserted at throat, 
stigmas located in distal part of tube or becoming slightly exserted. Capsules 
2-3 x 2-25 mm, usually slightly longer than wide, subglobose or broadly ob- 
long, slightly to somewhat compressed, thin- or rather thick-walled, fully infe- 
rior, apex rounded, retuse or truncate, tardily dehiscing loculicidally by a nar- 
row opening. Seeds ca 30-50 per capsule, (0.2-)0.3-0.4 mm diam., subglobose, 
ellipsoid, or ovoid, hilum punctiform, centric, testa reticulate, areoles polygonal, 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 321 


their walls thick, sinuous, testa surface verrucose. Flowering May to September. 
Chromosome number not known. 

Distribution and habitats—Mexico: endemic in Rio Verde area, southern 
part of San Luis Potosi. Fourteen collections have been seen. Rio Verde is some- 
times spelled Rioverde. Saline or alkaline flats or plains, sometimes in gypseous 
soil, associated with Prosopis and Acacia. 

Discussion.—B.L. Robinson cited Pringle 3758in the protologue, but did not 
designate the GH specimen as holotype. Robinson worked at the GH and we 
are designating the GH specimen as lectotype, from among at least nine 
ieaiccl eee 


D 


MEXICO. San Luis Potosi: 5.6 mi E of jct. to El Centro in Rio Verde on hwy 
70/86, mpio.Rio Verde, 21°54 N, 99° 53' W, Nesom 6635 (TEX); 0.7 mi N of Las Tablas (RR prrces 
mpio.Ciudad del Maiz, Nesom 6674 (TEX); Media Luna, near Rio Verde, Palmer 68 (CM, F GH, 
2 US); Minas de San Rafael, Guascama, Purpus 5014 (BM, F GH, MO, NY, US); San Bartolo, mpio. Rio 
erde, Soe ki 5821 (ENCB, MICH); 5 km S of Rio Verde, wae el camino a El Zapote, ae 
sa2e “B); 9 km E of Rio Verde, carretera a Rayon, Rzed 1 2478, cee RIZ, ENCB, MICH 
sa ee Las Tablas, Takaki 44 (ENCB); 2 km W of Estacion pene a, 30 km N of Rio Verde, 
akaki a (ENCB); 9 km SE of Villa Juarez, mpio. Villa Juarez, Takaki oe (ENCB). 


6. Oldenlandia salzmannii (DC.) Benth. & Hook. f. ex B.D. Jacks., Index Kew. 1:142. 
1893; 2:336. 1893. Anotis salzmannii DC., Prodr. 4:433. 1830. Hedyotis salzmannii (DC.) 
Steud., Nom. Bot. ed. 2.1:728.1840. TYPE: “circa Bahiam requens,” Salzmann s.n., nv 


Hedyotis thesiifolia A. St. Hil., Voy. Distr. Diam. 1:397. 1833. Oldenlandia thesiifolia (A. St. Hil.) K. 
Schum. in Mart., FI. Bras. 6, 6:270. P1127, £.1.1889.TYPE: n.v. 

Additional synonyms cited by Lewis (1966a:44). 
Small perennial herb, rooting at nodes and forming mats. Stems creeping, very 
slender, tetragonal-rounded, glabrous, often much branched. Leaves 1.5-5.2 x 
0.7-3 mm, rather thick, sessile or subsessile, broadly elliptic, elliptic, or ovate, 
glabrous to sparsely hirsute with pointed white hairs to 0.8 mm long, margins 
glabrous to ciliate especially at or near apex, base rounded, apex rounded. 
Stipules to ca 0.5 mm long, truncate, with O-few marginal fimbriae to ca 0.7 
mm long and O-few minute dark sessile glands. Inflorescence with flowers 
heterostylous, +-5-merous, solitary on slender pedicels 2-12 mm long. Hy- 
panthium densely to sparsely hirsute with widely spreading, white, pointed 
hairs ca’ 0.5-1 mm long; calyx lobes numbering 4-5(-6), 1-22 x 0.7-1.0 mm, 
lanceolate, ovate, or oblong, glabrous or glabrate, apices obtuse or rounded. Co- 
rollas 2.5-5 mm long, subsalverform, pink, lavender, light purple, or white, usu- 
ally with white center; tubes 1-2.2 mm long, ca 2 mm wide at base, broader 
than long, 2-3 mm wide at throat, tube and throat within densely hirsutulous 
or pubescent with spreading white hairs about 0.5 mm long; lobes numbering 
4(-5), 1-3.2 x 0.9-2 mm, as long as or longer than corolla tubes, ovate, spread- 
ing 90 degrees and apices recurved, striate-puberulent within with minute 
white hairs. Anthers numbering 4(-5), 0.5-0.8 x 0.2-0.4 mm, dark blue, oblong 


322 BRIT.ORG/SIDA 22(1) 


or elliptic. Stigma lobes 0.5-L.5 mm long, elliptic or linear, whitish, diverging at 
maturity. Pin flowers with anthers on filaments 0.3-0.7 mm long, attached in 
distal 1/3 of corolla tube; stigma lobes exserted 1-2 mm beyond throat, styles ca 
2-3mm long; slender, greenish or whitish. Thrum flowers with anthers exserted 
beyond throat on filaments 0.8-1.1 mm long; stigma lobes included in tube, style 
ca 1mm long. Capsules ca 1.5 mm long and wide, subglobose, hirsute with stiff 
white hairs, 7/8 to fully inferior, walls medium thick, dehiscence loculicidal. 
Seeds 4-14 per capsule, 0.3-0.5 mm long, black, trigonous, hilum apical, punc- 
tiform, testa reticulate, areoles polygonal or rectangular, areole walls distinct, 
sometimes double, testa surface apparently featureless. Flowering June to Au- 
gust in western Florida and adjacent Alabama. 

Chromosome number.—n = 15, 2n = 30. Lewis (1966b) as Hedyotis salzmannii, 
based on Lewis 6448 (MO), also in FTG, US! Pond edge in botanical garden, Rio 
de Janeiro, Guanabara State, Brazil. Lewis stated that meiosis was irregular and 
about 2/3 of pollen grains were shriveled. Terrell examined the US specimen 
and concluded that the plant does not closely resemble the plants from Florida. 
Thus, this chromosome number must be considered as tentative. 

Distribution and habitats.—South America: Brazil, Argentina, Chile, Uru- 
guay, Paraguay. United States: Locally adventive in Florida near Pensacola in 
Escambia and Santa Rosa counties and in adjacent Baldwin Co., Alabama, in 
roadside ditch, by ponds, and at edge of a marsh. First discovered by J. R. 
Burkhalter in Escambia Co. (Fosberg & Terrell 1985). Burkhalter sent Florida 
plants to Terrell, who grew them in a greenhouse for several years. 

Discussion.—The name, O. salzmannii, is here accepted as correct pending 
study of the type specimens. In Index Kewensis, vol. 1, B.D. Jackson, editor, listed 
“Anotis Salzmanni DC,, Le. =Oldenlandia Salzmanni.” In volume 2, p. 336, Jack- 
son listed “Oldenlandia Salzmanni Benth. & Hook. f. Gen. ii. 58.—Bras.” It was 
pointed out by Fosberg and Terrell (1985) that Bentham and Hooker did not 
validly publish the combination Oldenlandia salzmannii because they merely 
listed it under Oldenlandia, and did not state anything further about their in- 
tentions or viewpoints. Jackson validated the name Oldenlandia salzmannii 
by publishing it in volume | of Index Kewensis. We may alternatively cite only 
B.D. Jackson. 

The description of the later name Hedyotis thesiifolia A. St. Hilaire 1833) 
states that the leaves are linear, which casts some doubt on its identity, as the 
subject plants have ovate leaves. Later, Schumann (1889; see above) published a 
very good description of H. thesiifolia along with an excellent drawing, citing 
several synonyms, including Hedyotis salzmanni. Unfortunately, we have to 
stick with the earliest name, O. salzmannii, the one chosen by Fosberg & Terrell 
(1985). The taxonomy and nomenclature of this species need further study in- 
volving the variation in the native South American plants. 


—_— 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 323 


Collections from U.S.A. ALABAMA. Baldwin: Near Orange Beach, just N of W end of Cotton Bayou, E 
of Ala. 161, Burkhalter 8753 (UWFP). FLORIDA. Escambia: near Pensacola, S of US 98, 0.2 mi W of 
Fairfield Drive, Burkhalter 5408 (FLAS); Gonzalez, N of Chemstrand Road (FL C-297), E of Judy St. 
R30W, TIN, Sec. 14, Burkhalter & Hand 6537 (FLAS, FSU, UWFP); SW of Pensacola, E side of Herrion 
Bayou, N of US 98, Burkhalter 8838 (FLAS, UWFP). 


7. Oldenlandia eee ee & Cham.) DC,, Prodr. 4:428. 1830. re 
microtheca Schltdl. & , Linnaea 5:169. 1830. Hedyots fe ee (Schltdl. @ Cham 
Steud., Nomencl. Bot. ees TYPE: MEXICO. V Uz: “Barranca de erie in 
ee ae Sep 1828, Schiede & Deppe 390 one HAL); see further notes on 
isotypes in text below. 

Gerontogea deppeana Schltdl. & Cham., Linnaea 5:169. 1830. Oldenlandia deppeana (Schltdl. & 
Gon ) DC, Prodr. 4:428. 1830. Hedyotis oo (Schltdl. & Cham.) Steud., Nomencl. Bot. 
ed. 2. 1:727.1840.TYPE: MEXICO. VERACRUZ: San Andres, Aug 1828, Schiede & Deppe 391 (HO- 
LOTYPE: HAL} IsOTYPES: HAL! lca es 


Perennial herbs with roots sometimes thick and woody. Stem base of ten woody, 
to 5mm thick. Stems 9-42 cm tall, slender, erect or decumbent, glabrous, pu- 
berulent, or densely scabrid, branches to 30 cm long. Leaves subsessile or with 
partly winged petioles to 5 mm long, blades (6-)10-30(-40) x 2-13(-20) mm, l- 
nerved, quite variable in size and shape, ovate, lanceolate, or elliptic, varying to 
linear in Chiapas populations, glabrous to scaberulous or puberulent above, or 
densely scabrid in Chiapas populations, glabrous beneath, bases cuneate, 
rounded, or tapering, apices obtuse, acute, or subacuminate. Stipules toca 2 mm 
long, to3mm wide, scarious, whitish, deltate to lanceolate, glabrous or puberu- 
lent, with marginal gland-tipped teeth to ca 1 mm long, apices truncate to 
acuminate or lobed. Inflorescence terminal, in few-flowered cymes, flowers 
heterostylous, pedicels to 15 mm long, slender or filiform, glabrous or 
scaberulous. Hypanthium glabrous or with minute hairs on margins of sinuses; 
calyx lobes 0.5-13 x 0.3-0.5 mm, usually 1/4 as long to subequal to corolla 
tube, deltate to lanceolate, apices obtuse or acute. Corollas 2.5-7 mm long, fun- 
nelform, white or lined with purple, glabrous externally; tube 1-3 mm long, 
somewhat widened distally, 0.5-1.3 mm wide at base, 1-2 mm wide at throat, 
puberulent within with dark gland-tipped hairs 0.1-0.2 mm long: lobes 1.5-3(- 
4) x 0.8-18 mm, ovate or lanceolate, glabrous or puberulent within. Anthers 
0.4-1 mm long, 0.2-0.3 mm wide, elliptic or oblong. Stigma lobes to 1.4 mm 
long , linear, often somewhat coiled at maturity. Pin flowers with corolla lobes 
1-2(-3) times longer than corolla tube, anthers located just below corolla throat, 
sessile, stigma lobes exserted 1-2 mm beyond corolla throat, styles ca 2-3.2 mm 
long. Thrum flowers with corolla lobes 2/3-1 times as long as corolla tube, an- 
thers exserted beyond throat on filaments ca 0.3-1 mm long, stigma lobes lo- 
cated near midpoint of corolla tube. Capsules 1-3 x 1.5-3.5 mm, usually wider 
than long or subequal, subglobose to obovate, slightly to rather strongly com- 
pressed, glabrous or glabrate, 3/4-9/10 inferior, thin- or rather thick-walled, 


324 BRIT.ORG/SIDA 22(1) 


apices retuse, truncate, or broadly rounded, dehiscing loculicidally and then 
septicidally. Seeds 10-32 per capsule, 0.4-0.7 mm long, black or dark brown, 
irregularly and obtusely angulate or conoidal-angulate, hilum appearing ei- 
ther apical or centric, punctiform, testa not reticulate, areoles lacking, strongly 
coalesced, replaced by entangled vermiform strands. Flowering throughout the 
year. Chromosome number: n = 11 (Lewis in Terrell et al. 1986). 

Distribution and habitats —Mexico: Chiapas, Hidalgo, Oaxaca, Puebla, San 
Luis Potosi, Veracruz. Standley (1918) included Yucatan in the range, but I have 
not seen any specimens from there. Gravelly, rocky or sandy soil, slopes, banks, 
crevices of cliffs, in open or in pine or deciduous forests, and evergreen cloud 
forest in Chiapas, alt. 1000-2700 m. 

Discussion.—This species is closely related to O. ovata and O. drymarioides 
as shown by its morphology. Chiapas collections by Breedlove and Raven are 
linear-leaved and scabrid and occur in cloud forests. They were judged to be a 
possible new variety, but they appear to be otherwise similar to O. microtheca 
and, lacking more collections, are tentatively included in O. microtheca s.l. 

The type specimens for O. microtheca have been seen in the Halle (HAL) 
and other herbaria, but have presented problems in interpretation of label data. 
In addition to the holotype cited above at HAL, other collections included pos- 
sible isotypes, as follows: (1) a specimen at K labelled as 390, but lacking a local- 
ity; (2) a specimen at HAL labelled 390 and “Barranca de Tioselos,” but dated 
August 1829, not September 1828; (3) a type at B presumably destroyed, shown 
in photos of “Types of Berlin Herbarium” deposited at F MO, NY, US; this la- 
belled as “O. micrantha n. sp.” and as Hedyotis micrantha Schlecht. a name un- 
published in these combinations, (4) a specimen at HAL and MO, supposedly 
not a type, labelled from “In sylvis Papantlae,” thus agreeing with the citation 
under O. microtheca in Linnaea 6:414. 1831. The protologues of Gerontogea 
microtheca and G. deppeana appeared under the numbers 390 and 391 on the 
same page. Any question of priority seems to have been settled by Standley 
(1918), who adopted O. microtheca in his monographic treatment of 
Oldenlandia for the North American Flora. 

Standley (1918) listed O. latifolia Martens & Galeottii as a synonym of O. 
microtheca, but these are two separate species that are superficially so much 
alike, even in floral details, that it is difficult to identify them without the seeds. 
Like O. ovata and O. drymarioides, the seeds of O. microtheca are irregularly 
angulate or irregularly conoidal and have coalescent areoles as described here. 
In Hedyotis latifolia (Martens & Galeotii) Walpers, however, the seeds are flat 
and reticulate. The latter has a chromosome number of n = 17, compared to n 
=l1 in O. microtheca. A detailed study of the morphology of all parts of these 
two species in separate genera indicated that the best distinguishing charac- 
ters are those in the following key. 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 325 


KEY TO OLDENLANDIA MICROTHECA AND HEDYOTIS LATIFOLIA 


. Seeds 0.4-0.7 mm long, ices arly angulate or conoidal-angulate, testa not reticu- 
late, areoles appeari sced, replaced by entangled vermiform strands; hilum 
not or scarcely Poeible corolla tube 1-3 mm long, 1-2 mm wide at throat; corolla 
lobes in pin flowers 1-2(-3) times longer than tube, lobes in thrum flowers 2/3-1 
times as long as tube; anthers in pin flowers attached just below corolla throat 
Oldenlandia microtheca 


1. Seeds 0.5-1 mmlona flat, testa finel | ith | | : 
tiform, near center of ventral Gee of Cae corolla tube 1 5-5 mm long, 1.8-3 mm 
wide at throat; corolla lobes in both flower types 1/2-1 times as long as tube; 
anthers in pin flowers attached near midpoint of corolla tube Hedyotis latifolia 


Representative collections. MEXICO. Chiapas: Sumidero de Tuxtla, 22 km N of Tuxtla Gutierrez, 

00 ft., Raven & Breedlove 20123 (DS, NY); same loc., Breedlove 10650 (DS, NY); Summit of Chuchil 
Ton, NE of Bochil, mpio. of San Andres Larrainzar, 2700 m, Breedlove 29269 (DS, MICH, MO); 7 km 
NE of Bochil along road to Simojovel, mpio. of Bochil, Breedlove & Smith 32308 (DS); 11-15 km NW of 
Soyalo on road to La Bombana and Chicoasen, mpio. of Osumacinta, 1000 m, Breedlove 33756 (DS, 
MICH, MO); 6-8 km WNW of Soyalo, mpio. of Soyalo, pei 37168 (DS, ENCB, MICH, MO). Hidalgo: 
Vic. of Molango, road to Lolotla, Distr. Molango, Moore 2399 (GH, MICH). Oaxaca: Mpio. San Pedro 
Ixcatlan, Cerro Quemado, al W del Poblado de Cerro Quemado, Calzada 10351 (XAL). Puebla: 15 k 
adelante de Xicotepec de Juarez rumbo a Poza Rica, Chazaro B. 569 (XAL); Near Metlaltoyuca, ean. 
man 26 (GH, US); near Ocostoc below eee eas ised va TENN, US); Mpio. Cuetzalan, 
San Miguel Tzinacapan, Turra 1524 (ENCB); El Reparo, Mpio Ico, E ue 414 (CAS, 
CHAPA, ENCB, US); Carretera aoe San Andres, een Cuetzalan, Zola B. 253 (XAL). San Luis 
Potosi: Las Canoas, Pennell 17920 (F GH, MEXU, MICH, NY, PH, US); Tamasopo Canyon, Pringle 3510 
(FE GH, VT). Veracruz: Rancho La es a5 km al N de Banderilla, mpio. Banderilla, Calzada 
4327 (XAL): 1 km al E de San Pablo, mpio. de Papantla, Gutierrez R. 26 (ENCB); El Esquilon, mpio. 
Jilotepec, Ortega O. 75 (XAL); Monte Rey, Ejido Coetzalan, Mpio. Axocuapan, Robles H. 171 (ENCB, 
XAL); Below Txolo Falls near Jalapa, Sharp 45802 (GH, MEXU, TENN, US); 6 km SW of See 
mpio. Acatlan, road to Misantla, Terrell & Koch 5379 (CAS); Encinos, mpio. de Totutla, F Ventur 
7627 (CAS, ENCB); Coacoazintla, mpio. Coacoazintla, F Ventura A. 8292 (CAS, CHAPA, co 
Filipinas, mpio. de Tlapacoyan, E Ventura A. 12316 (CAS, CHAPA, ENCB). 


8. Oldenlandia ovata S. Watson, Proc. Amer. Acad. Arts 18:97. 1883. Hedyotis watsonii 
WH. Lewis, Rhodora 63:222. 1961 (non H. ovata Thunb. ex Maxim., Bull. Acad. Imp. Sci. Saint- 
Petersbourg 29: 161. 1883). TYPE: MEXICO. NUEVO LEON: Guajuco, Mar 1880, E. Palmer 399 (LEC- 
TOTYPE, here designated: GH; ISOLECTOTYPES: FI, K!, NA!, US!, VT!); LECTOTYPE mislabeled later 
as O. microtheca. PARATYPES: Ervend berg 200 (GH! atypical or not O. ovata); Parry and Palmer 
676 1/2 (GH); these in GH are on the same sheet as the lectotype. 


Small annual herb. Stems to ca. 20 cm tall, slender, weak, ascending, decum- 
bent, or diffusely spreading, glabrous or puberulent. Leaves with petioles to ca 
5mm long, blades 3-14 x 1-9.5 mm, thin, l-nerved, ovate, broadly ovate, or up- 
per leaves elliptic, glabrous to sparsely pubescent above, glabrous beneath, bases 
rounded, subcordate, or subtruncate or upper leaves tapering to base, apices 
obtuse or acute. Stipules ca. 0.5 mm long, truncate, with gland-tipped teeth. 
Inflorescence terminal and axillary in groups of 1-3 flowers, cymose, pedicels 
filiform, 2-16 mm long, flowers isostylous. Hypanthium glabrous; calyx lobes 
0.4-1.2 mm long, deltate to lanceolate, obtuse or acutish. Corollas 2-4 mm long, 


326 BRIT.ORG/SIDA 22(1) 


short-funnelform, white, glabrous or puberulent within; tubes 1-2.2 mm long, 
scarcely wider distally; lobes 1-1.5 mm long, ovate. Anthers 0.4-0.7 mm long, 
sessile, elliptic or narrowly elliptic, attached at corolla sinuses, slightly exserted. 
Stigma lobes 1-1.2 mm long, linear, slightly exserted at throat, not exceeding 
corolla lobes, style 1.3-1.7 mm long. Capsules 1.3-2.2 x 2-3 mm, subglobose or 
broadly oblong, slightly wider than long, usually widest at apex, glabrous, thin- 
walled, fragile, 3/4-4/5 inferior, subtruncate at apex, dehiscing loculicidally 
and then septicidally. Seeds ca. 17-34 per capsule, 0.4-0.7 mm long, black or 
brown, irregularly and obtusely angulate or conoidal-angulate, hilum scarcely 
visible, apical, punctiform, surface often warty and bumpy, testa not reticulate, 
areoles lacking, strongly coalesced, replaced by entangled vermiform strands. 
Flowering March to December. Chromosome number: 2n = 12 (Lewis 1962 as 
Hedyotis watsonii). 

Distribution and habitats.-MEXICO: Central and southern Nuevo Leon 
and Tamaulipas south to eastern San Luis Potosi, northern Hidalgo, and Ver- 
acruz (one collection). Moist shaded rocks and outcrops along streams, rock crev- 
ices, and on canyon walls, alt. 100-1500 m, 300-5000 feet. 

Discussion.—Additional nomenclatural data are provided as follows. The 
lectotype is labelled “March 1880,” but isolectotype F is labelled “March 1-8, 
1880” and states that Guajuco is 27 miles southeast of Monterrey; K and US are 
labelled “February to October 1880.” A paratype listed above is L.C. Ervend berg 
200 (GH), Wartenberg, near Tantoyuca, prov. Huasteca, 1858, en route from San 
Luis Potosi to Tampico, December 1878 to February 1879. Ervend berg 200 is ei- 
ther atypical O. ovata or not O. ovata; it is difficult to know which because of 
the condition of the specimen. 


eae: collections. MEXICO. Hidalgo: Near Chapulhuacan, Sharp 441822 (GH, MEXU, TENN, 
US). Nuevo Leon: Canyon Denuncio, Rancho La Bolla, Mpio. de ie a Santi: Haas Muel ler 2020 (F GH, 
MICH); Sierra Madre Orientale 21 mi W of Linares, Ripley & Barneby 13559 (NY); 0.5 mi SE of Canoas, 
5.5 mi SE of Allende, E. & B. Terrell 4426 (CAS): Trail below base of ey ig ee Monterrey, E. 
& B. Terrell 4430 (CAS); Hacienda Vista Hermosa, 35 mi S of Monterrey, White 1555 (MICH): Queretaro: 
4 km SW of Ahuacatlan, mpio. de Pinal de Amoles, Rzedowski 27744 (ENCB); San Luis Potosi: 
Roeaune it Fisher 45180 (BRIT, US). Tamaulipas: 22 mi SW of Victoria, Crutchfield & Johnston 

B( TEX); 3 mi us of Villagran on Victoria-Linares hwy., yeh nston & Graham 4672B (TEX); Mpio. 
Sener 43m of Adelaida-Guayabas jet., Nesom et al. 6351 (TEX); Cerro del Picacho, mpio. de 
Villa de Casas, ae 4098 (ENCB); ee ies Sabinas, opposite i 619 on Pan American hwy. N of El 
Limon, Sharp & Hernandez X. 50393 


9. Oldenlandia drymarioides (Standl.) Terrell, Phytologia 59:80.1985. Houstonia 
drymarioides Standl., J. Wash. Acad. Sci. 18:162.1928. Hedyotis drymarioides (Standl.) WH 
Lewis, Rhodora 63:221.1961.T ype: MEXICO. TAMAULIPAS: mountains S of Victoria, alt. LOOO m, 
9 Apr 1926, Robert Runyon 870 (HOLOTYPE, designated by Standley, US!: ISOTYPE: Fl). PARATYPE: 
same loc. and date, Robert Runyon & B.C. Tharp 4039 (TEX!, US). 


Annual herb (protologue states perennial, but plants appear annual). Stems to 
cal5cm tall, slender, erect or decumbent, glabrous, simple or sparsely branched, 


TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 327 


with the 3-4 internodes much longer than the leaves. Basal leaves to 12 mm 
long, to5 mm wide, much larger than cauline leaves, forming a rosette, ovate or 
oval, rounded at base, with short petioles, glabrous or minutely puberulent: 
cauline leaves to 6 mm long, to 3 mm wide, few and much reduced, elliptic to 
narrowly elliptic. Stipules 0.5 mm long, with short marginal teeth. Inflorescence 
in few-flowered, terminal cymes, flowers apparently isostylous, pedicels to ca 
18 mm long, filiform. Hypanthium glabrous; calyx lobes 0.5-1.2 mm long, equal- 
ling or slightly exceeding mature capsules, deltate. Corollas 2-4 mm long: short- 
funnelform, apparently white; tubes 1.3-2.3 mm long, scarcely wider distally: 
lobes ca 1-1.8 mm long, ovate. Anthers ca 0.8 mm long, sessile, oblong, attached 
at sinuses of corolla lobes, slightly exserted. Stigma lobes not seen Capsules 
13-2 x L7-28mm, slightly wider than long, slightly I d 
walled, fragile, 1/2-3/4 inferior, broadly rounded at apex, dehiscing 
loculicidally and then septicidally. Seeds 0.3-0.45 mm long, black, obtusely and 
irregularly angulate or conoidal-angulate, hilum scarcely visible, apical, punc- 
tiform, testa not reticulate, areoles lacking, strongly coalesced, replaced by en- 
tangled vermiform strands. Chromosome number not known. 

Discussion.—This description is shortened due to the few collections. Fur- 
ther collections are needed to determine whether this species is distinct from 
O. ovata. 


glabrous, thin- 


Recent collection. MEXICO: Tamaulipas: Mpio. San Carlos, Sierra de San Carlos, ca. 5 mi S of San 
Carlos, N side of Bufa El Diente, igneous bedrock, N-facing steep slope; scattered oaks and shrubby 
vegetation of Rhus, Croton, Chiococca, Helietta, Vauquilinia, 770 to 1100 m, 24°31.5' N, 98° 57.6’ W. 
Flowers white, very common on moist shady bank, 17 Jun 1987, Nesom 6049, with Norris, Martinez, 
Woodruff (MEXU, TEX). 


ACKNOWLEDGMENTS 


We thank Scott Whittaker and Susann Braden, SEM Laboratory, Smithsonian 
Institution, for preparation of illustrations, and Marjorie Knowles for format- 
ting the illustrations. We express appreciation to the curators of the herbaria 
cited for many loans of specimens or access to collections. Richard Wunderlin 
and Bruce Hansen provided facilities at University of South Florida where some 
work on this manuscript was carried out. Joseph Kirkbride and John Wiersema 
provided helpful discussions concerning difficult nomenclatural problems. 
Steven Ginzbarg, Collections Manager, UNA, assisted in selection of a lectotype 
for Oldenlandia littoralis. 
REFERENCES 
Anpersson, L.and J.H.E. Rova. 1999. The rps16 intron and the phylogeny of the Rubioideae 
(Rubiaceae). Pl. Syst. Evol. 214:161-186. 
ANDERSSON, L., J.H.E. Rova, and F.G. Atzate.2002.Relationships, circumscription, and biogeog- 
raphy of Arcytophylium (Rubiaceae) based on evidence from cpDNA. Brittonia 54: 


328 BRIT.ORG/SIDA 22(1 


= 


40-49. 

Bremekamp, C.E.B. 1952. The African species of Oldenlandia L. sensu Hiern et K. Schumann. 
Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 48:1-297. 

Bremer, B. 1996. Phylogenetic studies within Rubiaceae and relationships to other families 
based on molecular data. Opera Bot. Belg. 7:33-50. 

Bremer, B.and J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae 
(Rubiaceae). PI. Syst. Evol. 225:43-72. 

FosperG, F.R.1943. The Polynesian species of Hedyotis 
1-102. 

Fospera, F.R.and E.E. Terrett. 1985. A recently established exotic in west Florida and Alabama 
(Hedyotis salzmannii or Oldenlandia salzmannii; Rubiaceae). Castanea 50:49-51. 

Hate,N.1966. No. 12, Rubiacées (1° part), Hedyotidées. In: Aubréville, A., editor. Flore du 
Gabon. Mus. Hist. Nat., Paris. Pp. 75-124. 

Hepper, FN. and R.W.J. Keay. 1963. Rubiaceae (part), vol. 2, Oldenlandia. In: FN. Hepper, ed. 
Flora of West Tropical Africa. 2nd ed. Crown Agents, London. Pp. 210-212. 

HitcHcock, A.S.and M.L.Gareen. 1929. Standard-species of Linnaean genera of Phanerogamae. 
Intl. Bot. Congress, Cambridge, 1930. Nomencl. Prop. Brit. Bot. 110-199. London. 

Howarb, R.A. 1989. Flora of the Lesser Antilles. Vol. 6, Part 3. Arnold Arboretum. 

Jarvis, C.E., FR. Barrie, D.M. ALLAN, and J.L. Reveat. 1993. A list of Linnaean generic names and 
their types. Koeltz, KOnigstein 

Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. 
Bot. 49:855-865. 

Lewis, W.H. 1964. Oldenlandia corymbosa (Rubiaceae). Grana Palynolog. 5:330-341. 

Lewis, W.H. 1965. Cytopalynological studies of African Hedyotideae (Rubiaceae). Ann. Mis- 
souri Bot. Gard. 52:182-211. 

Lewis, WH. 1966a. The Asian genus Neanotis nomen novum (Anotis) and allied taxa in the 
Americas (Rubiaceae). Ann. Missouri Bot. Gard. 53:32-46. 

Lewis, WH. 1966b Chromosome numbers of Phanerogams. 1. Ann. Missouri Bot. Gard. 
53:100-103. 

Lewis, WH. 1966c. Chromosome numbers of Oldenlandia corymbosa (Rubiaceae) from 
southeastern Asia. Ann. Missouri Bot. Gard. 53:25 7-264. 

Sivarasan, VV. and S.D. Buu. 1990. Taxonomic and nomenclatural notes on the Hedyotis 
corymbosa-diffusa complex (Rubiaceae) in India. Taxon 39:665-674. 

STANDLEY, P. 1918. Rubiaceae, Oldenlandieae, in, North American flora 32, part 1:17-39. 

Terrett, E.E.1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 
68:125-133. 

TerreLt, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48. 
|- : 

Terrett, E.E., W.H.Lewis, H-Rosinson, and J.W.Nowicke.1986.Phylogenetic implications of di- 
verse seed types, chromosome numbers, and pollen morphology in Houstonia 
(Rubiaceae). Amer. J. Bot. 73:103-115. 


— 


Rubiaceae). Bishop Mus. Bull. 174: 


TERRELL AND ROBINSON, TARAVINUIVIE UE 329 


Terrett, E.E. and H. Rosinson. 2003. Survey of Asian and Pacific species of Hedyotis and 
Oldenlandia (Rubiaceae) with nomenclatural notes on Hedyotis types. Taxon 52: 


Terrett, E.E.and R.P. Wunber.in. 2002. Seed and fruit characters in selected Spermacoceae 
and comparison with Hedyotideae (Rubiaceae). Sida 20:549-557. 

Verdcourt, B.1976. Rubiaceae (Part 1), Oldenlandia.|n:R.M. Polhill, ed. Flora of Tropical East 
Africa. Crown Agents, London. Pp. 268-315. 

Verdcourt, B.1989. Rubiaceae, Oldenlandia. In: Flora Zambesiaca, vol. 5, part 1. Flora 
Zambesiaca Managing Committee, London. Pp. 120-142. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Ricarpo AyerzA Jk.and Wayne Coates. 2005. Chia: Rediscovering a Forgotten Crop 
of the Aztecs. (ISBN 0-8165-2488-2, pbk.). University of Arizona Press, Tuscon, 
AZ 85719, US.A. (Orders: http://ww ae ona.edu/). $14.95, 215 pp., 
b&w enna tables, index, 6" x 


The plant | Salvi hispanica and Salvidcol } i \} 


part of the human diet, 
medicine chest, and culture for 4000 years. Although its’ importance to humans has waned over 


time, there is currently a renewed interest in chia both as a food and a good source of omega-3 fatty 


acids. Authors Ayerza Jr and Coates have created an interesting primer on the history and uses of 
chia, as well as the pabake values of the species. 


luce the reader to th lithi learl so-American diets and 
compare oe toa ace western diet. Although these early diets were ‘high in animal products 
like ours today, theirs relied heavily on lean game animals from the wild; ours consists mostly of 
grain fed domesticated animals. This comparison of diets and the introduction of omega-3 and omega- 
6 fatty acid ratio prepare the reader for a detailed discussion of fats in the human diet. 

uthors have written the text on fats and oils in the human diet ina very readable way. The 
information is well referenced with research and related literature. Each type of lipids, cholesterol 
and fatty acids is discussed. Information is presented on coronary heart disease as well as the detri- 
mental increase of omega-6 fatty acids and decrease of more healthful omega-3 fatty acids in the 
western cliet. 


After this in-depth discussion of how the omega-3/6 fatty acids fit into our diet, the authors 
delve into chia even further. The text includes a chapter on the history and uses of chia through 
information collected from codices, written histories and related cultural uses of today. Chia was 
important in the Aztec empire as a food, beverage, medicine, soup, and religious ceremonies. The 
S] anish ¢ nquest caused a decline in the use of chia because the invaders banned chia to help deter 


non-Christian religious ceremonies. After so many « uries, Cl s still gr inM 1 Soutl 


America, but i wnotas impor rfantac rop acir once was. 


12 : is ] ee 


The remaining chapters of the | 


vays to determine 
the different specs botanical description, ecology, and cuevation information on the chia pene 
Rel ated tO the earliet discussion of human die SL 


as a good source of omega-3 fatty acids. They also compare it to other omega- ge sources and discuss 

the potential use of chia in enriching other foods sources such as eggs with omega-3 fatty acids. One 

very interetne section n included an introduction to the Nahua system of botanical classification. 
CI 


a Forgotten Crop of the ii tecsisa poo filled “ ae anaes 
about chia, its’ s history. uses, and nutritive value. 
] 


ifot mative 
as a functional food with ee nutrient ae substantial amounts of vita- 
mins and miner a ow sodium content, a substantial source of omega-3 fatty acids, and are a non- 
allergenic food with antioxidants and fiber. This book can be recommended to anyone interested in 
learning more about omega-3 fatty acids, and chia’s nutritive value or history.—Lee Luckeydoo, Her- 
barium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(1): 330. 2006 


NEW TAXA AND COMBINATIONS IN CULTIVATED 
BAMBOOS (POACEAE: BAMBUSOIDEAE) 


C.M.A. Stapleton 
Royal Botanic Gardens, Kew 
Richmond, Surrey 

UK 


chris_stapleton@onetel.com 


ABSTRACT 


A new species in Fargesia Franch. from China, and a new variety in Drepanostachyum falcatum (Nees) 
Keng f. from the eastern Himalayas, are described. Foun: new Son ee are Eade in Borinda 
Stapleton for species from western China, previously irge new com 


bination is made in Indocalamus Nakai for a Japanese species deeaed: in Bune Nakai 
RESUMEN 


te J st * ] om ‘ | 4 


h. de China, y una nueva variedad de Drepanostachyum 
faxed (Nees) Keng f. ‘del Este del Himalaya. Se hacen crane combinaciones nuevas en Borinda 


Stapleton para especies del Oeste de China, p en el género Fargesia, y se hace 


una combinacion nueva en Indocalamus Nakai para una especie japonesa descrita en el género 
Pseudosasa Nakai. 


Fargesia apicirubens Stapleton, sp. nov. Type: UK: Ness B ic Garden, C.M.A. Stapleton 
1060, 11 May 1996 (HOLOTYPE: K!). 


Fargesia robusta foliis crassis lanceolatis vel 
vaginorum culmorum glabris differt. 


Culms 2-4 m, 0.3-1 cm in diam., pendulous; internodes 5-18 cm, smooth, lightly 
white-powdery initially, becoming glossy dark green with purple spots, often 
apically purple-red, later orange-yellow, wall to 5 mm thick; nodes with weakly 
prominent supra-nodal ridge, sheath scar level and tall, mid-culm branches 6-8. 
Culm sheaths slowly deciduous, basally leathery and smooth, light brown with 
prominent red-brown ribs, apically asymmetrically rounded with a persistent 
red band below the blade, much shorter than internodes, glabrous, margins 
densely yellow-brown-setose; auricles absent; oral setae absent; ligule ca. 1 mm, 
truncate, initially ciliolate; blade lanceolate, erect or reflexed, glabrous, margins 
setose. Leaves 3-7 per ultimate branch; sheath thick, glabrous with outer margin 
densely white-ciliate at first, usually apically red with yellow veins; auricles and 
oral setae absent ligule ca. 1 mm, truncate, purple, shortly tomentose, external 
ligule not evident, tomentose; blade lanceolate to fusiform, 7-10 = 0.6-1.5 cm, 
slightly thickened, glabrous, basally cuneate, apically long-acuminate, secondary 
veins 4-5-paired, one margin shortly spinescent, the other obscurely serrulate, 
transverse veins distinct, petiole yellow. Inflorescence racemose, unilateral; spike- 
lets shortly pedicellate, usually subtended by densely pubescent reduced sheaths. 

Introduced from Daba Shan, Shaanxi Province, China, and grown under 
the misapplied name Fargesia dracocephala TP. Yi. 


SIDA 22(1): 331 — 332. 2006 


332 BRIT.ORG/SIDA 22(1) 


Etymology.—The epithet “apicirubens” refers to the distally red-colored 
sheaths and culm internodes. 


sd aaa Oia falcatum var. sengteeanum Stapleton, var. nov. Type: U.S.A. OR- 
N: Gold Beach, Aug 1996, S.G. Cooper s.n. (HOLOTYPE: K!). 


=— 


Drepa ie ee THK Keng f. var. falcato spiculis bifloris vel trifloris, culmis porcatis, foliis 
angustis similis, sed vaginis culmorum praeter annulum pilosum circumcirca fundum omnino 
slabris, lemmatibus scabris, paleis vix bifidis, rhachillis minus puberulis, pilis costae foliarum 
longioribus unifariis dispersis, vaginis foliorum haud cilatis differt. 
Culms 2-4 m, 4-10 mm in diam, internodes 10-21 cm, uniformly finely porcate, 
lightly white-waxy at first; nodes substantially raised, with ring of persistent, 
initially orange-brown, later colorless, 1-3 mm hairs. Culm sheaths abaxially gla- 
brous except for a dense ring of 1-3 mm red-brown hairs around basal 1-2 mm; 
margins distally with ca. 1mm white cilia; auricles absent; oral setae absent; ligule 
long-fimbriate, abaxially basally puberulous and apically glabrous. Leaf sheaths 
glabrous; external ligule pronounced, thin, with even ca. 0.1 mm cilia; auricles 
glab ,elevated; oral setae absent; ligule ca. 1mm, rounded to 
acute, shortly laciniate, shortly ciliate, basally puberulous. Leaf blade to 12 x | 
cm, basally narrowly cuneate; abaxially glabrous except for sparse ca. | mm hairs 
on one side of the proximal midrib; adaxially glabrous. Spikelets with (1-)2-3(- 
4) florets. Lemma 7-10 mm, deeply furrowed, abaxially distally scabrous, distal 
margins shortly ciliate. Palea 8-10 mm, abaxially distally shortly scabrous, apex 
acute or very shortly bifid, shortly ciliate; keels distally scabrous. Rhachilla sec- 
tions to5 mm, mainly glabrous, apically bearded with ring of ca. 1 mm hairs. 
Cultivated in U.S.A. and Europe under misapplied names that will be dis- 
cussed in a further article. Presumably originating from the Himalayas. 
Etymology.—This taxon is named in honor of Dr. Lee Seng Tee. 


absent, 


Selected specimens. UK (cult.): Kew, Temperate House, 1894, Anonymous (K); Kew, Sep 1895, Anony- 
ous (K); Kew, Temperate House, 5 Jan 1897, Gamble s.n. (x); Kew, oe louse, 14 Sep 1901, 
oe (K). U.S.A. California: Bonny Doon, Stapleton 1136, 6 Oct 1999 (kK 


— angustissima (T.P. Yi) aes comb. nov. BasionyM: Fargesia angustissima 
Yi, J. Bamboo Res. 4(2):21. 198 
Borinda contracta (TP. bee saplton comb. nov. BasIoNyM: Fargesia contracta TP. Yi, J. 
mboo Res., 7(2):60 
Borinda nujiangensis ae & C.M. Hui) Stapleton, comb. nov. Basiony™: Fargesia 
nujiangensis Hsueh @ C.M. Hui, Bull. Bot. Res., Harbin 18:261. 1998. 
Borinda utilis or Stapleton, comb. nov. BASIONYM: Fargesia utilis T.P. Yi, J. Bamboo 
Res. 7(2):28 


a hamadae ou ) Stapleton, comb. nov. Basionym: Pseudosasa hamadae 
atus., J. Geobot. 15:86. 196 
ACKNOWLEDGMENTS 


[thank Mary Barkworth (UTC) and Kanchi Gandhi (GH) for promptly review- 
ing this note. 


ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XXIV: 
BAHIELLA (APOCYNOIDEAE: ECHITEAE), 
UN DESAPERCIBIDO NUEVO GENERO 
ENDEMICO DE BAHIA, BRASIL 


J.Francisco Morales 


Instituto Nacional de Biodiversidad (INBio) 
pto 22-3100 
Santo Domingo de Heredia, COSTA RICA 


RESUMEN 
BahiellaJ.F Morales (Apocynoideae Echiteae) 6 dé ] lo de Bahi Brasil 
es descrito, proponiéndose una nueva combinacion, B. Blanchet (A. DC.) J.F Morales y una nueva 
especie, B. infundibuliflora J.F Morales. Se incluyen descripciones completas, ilustraciones y 
Specunenes eapouinages Seen ee se realiz6 un analisis cladistico para determinar las 
la, el cual incluy6 un total de 44 especies (19 géneros) de 
i tribu Echiteae como Bae interno y 10 especies (7 géneros) de las tribus Apocyneae, Malouetieae 
y Mess como pen unOss externos. Hees resultados sugieren que Bahiella pertenece a las Echiteae, 

Ee 


p g ser determinadas por ahora. 


ABSTRACT 


] - oy R a ee pa a P| 4 


Bahiella J.F Morales (Apocynoideae, Echiteae), a new g 
illustrated. A new Reconstr B. hana: (A. Peel Morals: sane a new species, B. 
infundibulif d ] Aditionally. 


a iL 
13-14 : ] : ft if J . 1} cg 2 : ] : Ks ft 


es 44 species a9 genera) as ingroup of the tribe Echiteae, aod 10 species (7 ae of the 
ribes Apocyneae, Malouetieae y Mesechiteae as outgroup. The results suggests that Bahiella belong 
to the Echiteae, but their intergeneric relationships can’t be established by now. 


Desde su gecaion os Pale de eu (1810), la familia Asclepidaceae fue 
tradici if tedelas A Apocynaceae, pero estudios 
moleculares en la tltima década (e.g., Judd et al. 1994; Potgieter @ Albert 2001; 
Sennblad & Bremen 1996) sugirieron que ambas debian tratarse como una sola 
unidad. En forma reciente, Endress & Bruyns (2000), propusieron un nuevo 
sistema de clasificacion tratando en forma conjunta los géneros anteriormente 
divididos en Apocynaceae y Asclepiadaceae, reconociendo un total de cinco 
subfamilias. De estas, la subfamilia Apocynoideae esta dividida en cinco tribus, 
Apocyneae, Echiteae, Malouetieae, Mesechiteae y Wrightieae, separadas 
principalmente por las caracteristicas de la cabeza estigmatica. Sin embargo, la 
conformacion definitiva de estas tribus atin se encuentra en disputa, dado que 
analisis moleculares recientes (e.g., Simoes et al. 2004) sugieren que al menos 
una de ellas no es monofilética (Apocyneae) y mas estudios son necesarios con 


SIDA 22(1): 333 - 353. 2006 


334 BRIT.ORG/SIDA 22(1) 


el objeto de definir apropiadamente sus limites. Ahora bien, la mayoria de 
géneros neotropicales de las tribus Echiteae y Mesechiteae fueron 
monografiados por Robert Woodson (1933, 1935, 1936), siendo amos grupos 
relativamente faciles de distinguir por la ausencia 0 presencia de coléteres en la 
superficie adaxial del nervio central, asi como por las caracteristicas de la cabeza 
estigmatica. En los miembros de Mesechiteae, las laminas foliares tienen 
coléteres en la superficie adaxial del nervio central, ya sea agrupados en la base 
o distribuidos en forma irregular a lo largo de su entera longitud, aunque 
también pueden estar } en las venas secundarias (i.e., Mandevilla) y las 
cabezas estigmaticas tienen cinco costillas o proyecciones longitudinales, las 
cuales pueden estar restringidas a la base o distribuidas a lo largo de la cabeza. 
En contraposicion, los miembros de la Echiteae, tienen laminas foliares sin 
coléteres en la superficie adaxial de los nervios, asi como cabezas estigmaticas 
sin costillas longitudinales y con forma de carrete de hilo o mas o menos 
fusiformes y con un anillo membranoso basal. 

Dado que las monografias publicadas por Woodson se encuentran en su 
mayoria desactualizadas, se ha iniciado un proceso paulatino para hacer nuevas 
revisiones de los géneros de las tribus Echiteae y Mesechiteae que no hayan sido 
monografiados en los Ultimos anos. Por el momento, esto ha producido la 
revision total o parcial de varios grupos (e.g., Hansen 1985; Morales 1997 c, 1999 
b, 2002 a, 2002 b, 2003; Williams 1998), la descripcion de nuevos géneros (Mo- 
rales 1999 a; Morales & Williams 2004; Williams 2002) y la publicacion de 
varias nuevas especies. En el proceso de revision del género Prestonia, se procedié 
a examinar la mayoria de colecciones tipos disponibles, tanto de especies 
descritas en ese género, como de aquellas ubicadas en otros diferentes (e.g., 
Echites), pero que han sido sinonimizadas a través del tiempo en Prestonia. 
Echites blanchetii fue descrito por Alphonse de Candolle (1844), basadoen una 
coleccion hecha por Blanchet (3223 a) en 1840 en el estado de Bahia, Brasil. Este 
nombre fue reducido a la sinonimia de Prestonia coalita (Vell.) Woodson por 
Woodson (1936), quien probablemente nunca vio el tipo, ya que no cité dicha 
coleccion entre la lista de especimenes examinados. Luego de concluir la 
monogratia de Prestonia, era claro que Echites blanchetii fue incorrectamente 
sinonimizado bajo P. coalita y que ademas no podia ser ubicado incluso en ese 
género, pues sus corolas carecian de estructuras coronales accesorias (i.e., Co- 
rona anular alrededor de la fauce 0 corona epiestaminal, compuesta por cinco 
I6bulos coronales libres en forma opuesta a cada antera), uno de los principales 
caracteres diagnosticos de Prestonia. Ahora bien, mientras trabajaba en la 
elaboracion de una nueva monografia para los géneros Mandevilla y Mesechites, 
noté que varias colecciones provenientes del estado de Bahia en Brasil, 
identificadas a través de los anos como M. mansoanus (A. DC.) Woodson y 
morfoldgicamente algo similares a esa especie, tanto en la forma de las hojas, 
comoen la estructura de la inflorescencia, tenian hojas eglandulares, careciendo 


MORALES, BAHIELLA F f ECTAN oe 


de coléteres en la superficie adaxial del nervio central. Aunque la presencia de 
coléteres en el nervio central es algo comtin para varios géneros de la tribu 
Mesechiteae, muy raramente algunas especies pueden carecer de coléteres (e.g, 
Mandevilla angustifolia (Malme) Woodson). Sin embargo, al proceder con el 
analisis de las flores, pude descubrir que la cabeza estigmatica de este grupo de 
especimenes tiene forma de carrete de hilo, mas angosta medialmente y con un 
anillo membranoso basal, sin costillas o crestas basales longitudinales, esto 
ultimo un caracter diagnéstico que poseen el resto de miembros de la tribu 
Mesechiteae, a la cual pertenece Mesechites (Simoes et al. 2004). Este material 
coincide en todos sus caracteres morfol6gicos con el tipo de E. blanchetii, asi 
como a otras escasas colecciones depositadas en diferentes herbarios europeos. 
De esta manera, siguiendo a Endress & Bruyns (2000) y tomando el cuenta las 
caracteristicas de la cabeza estigmatica, est pertenecen a latribu 
Echiteae, pero no estan relacionados con ninguno de los géneros hasta ahora 
conocidos dentro de esa tribu. Por lo tanto, un nuevo género se propone a 
continuacion, incluida una descripcion de sus caracteres morfologicos, con 
discusidn de sus relaciones intergenéricas, una nueva combinacion y la 
descripcion de una nueva especie. 


MORFOLOGIA 


Habito y tallos 

Las dos especies de Bahiella son lianas, con los tallos viejos conspicuamente 
lefosos. En forma general, las ramitas son algo aplanadas bajos los nudos cuando 
jovenes, volviéndose teretes a subteretes con la edad. En algunos especimenes y 
en forma mas rara, tallos jovenes levemente angulados son encontrados. La 
secrecion de las ramitas y otras partes de la planta al cortarse es blanca. En 
contraposicion con especies de géneros de la tribu Mesechiteae (e.g., Mandevilla, 
Mesechites), donde los coléteres de los nudos pueden ayudarnos a reconocer 
ciertos grupos de especies, en Bahiella (asi como en el resto de los géneros de la 
tribu Echiteae) los coléteres intrapeciolares son diminutos, fusiformes e 
inconspicuos y no tienen utilidad para distincion especifica a nivel genérico. 


Hojas 

Las hojas son siempre pecioladas, no peltadas y eglandulares, sin coléteres en el 
nervio central adaxialmente, con la lamina conspicuamente coridcea, 
basicamente glabra y los margenes conspicuamente revolutos marginalmente. 
En los demas miembros de la tribu Echiteae, es muy raro encontrar especies 
con los margenes revolutos, estando limitados principalmente a Hylaea y 
ocasionalmente y en forma muy rara a unas pocas especies del género Prestonia 
y Echites. En forma general, solo la vena principal esta impresa en ambas su- 
perficies, siendo las venas secundarias y terciarias usualmente no evidentes y 
no impresas. 


336 BRIT.ORG/SIDA 22(1) 


Inflorescencias 
Las inflorescencias son cimosas, axilares, ramificadas, con numerosas flores 


densamente aglomeradas en las partes distales de la inflorescencia, 


diminutamente puberulentas a glabrescentes, pedunculadas, con los pedicelos 
siempre conspicuos y evidentes. Las bracteas florales son siempre escariosas e 
inconspicuas y las bracteolas son ausentes. La presencia de bracteolas no es algo 
comun en las Echiteae y esta limitadoa unos pocos géneros (e.g., Macropharynx, 
Peltastes). 


Flores 
El caliz esta compuesto por cinco sépalos escariosos e inconspicuos, con el apice 
erecto o suberecto (no reflexo), los cuales albergan en la base de la cara adaxial 
un coléter solitario, irregular y profundamente lacerado apicalmente. En Bahiella 
blanchetii, los coléteres estan algunas veces profundamente coco por lo que 
erroneamente pueden ser tomados como una linea de colé como 
los presentes en el género Peltastes. Sin embargo, el grado de laceracion de los 
coléteres no parece tener ninguna utilidad taxonomica, dado el alto nivel de 
variacion intraespecifica presente en varios géneros de las Echiteae. 

La forma de la corola varia de hipocrateriforma a infundibuliforma, glabra 
a inconspicuamente puberulenta exteriormente, sin estructuras coronales 
interiores (en forma de corona anular alrededor de la fauce o de una corona 
epiestaminal conformada por cinco lobulos coronales libres en forma opuesta 
a cada antera). El limbo esta conformado por cinco lobulos, usualmente 
extendidos en la antesis, pero algo reflexos marginalmente, con estivacion 
dextrosa. Los estambres estan ubicados en la mitad distal del tubo y son 
totalmente incluidos, con las anteras glabras dorsalmente, fuertemente 
aglutinadas a la cabeza estigmatica en dos puntos, con sus bases estériles y 
sagitadas, con cada auricula conspicuamente acuminada 

El ovario es apocarpico y esta conformado por dos carpelos, glabros y unidos 
apicalmente al estilo, cada carpelo con numerosos vulos. La cabeza estigmatica 
tiene forma de carrete de hilo, relativamente fusiforme, con un conspicuo anillo 
membranoso en la base, siendo mucho mas ancha en su parte basal. El nectario 
es anular, pero con el borde superior irregularmente pentalobulado. La 
lobulacion del nectario puede ser muy leve y casi no evidente o estar dividido 
en lobulos relativamente bien marcados, incluso en el mismo especimen. Este 
tipo de variacion intraespecifica no es algo raro en las Echiteae, ya que tal y 
como se ha comentado en forma previa (Morales 2004), el grado de lobulacion 
del nectario puede ser muy alto es algunos géneros y por lo tanto, es un caracter 
que debe utilizarse con reserva a nivel de distinci6n interespecifico. 


Frutos y semillas 
Los frutos estan compuestos por dos foliculos, relativamente divaricados, 
continuos y glabros, libres, continuos, secos y levemente lenosos al secar, con 


MORALES, BAHIELLA 3 337 


diminutas costillas 0 protuberancias longitudinales. Foliculos divaricados 
tampoco son comunes en las Echiteae, estando presentes en unos pocos 
miembros de unos cuantos géneros (e.g., Echites, Prestonia, Stipecoma). Las 
semillas son numerosas, secas, glabras, comosas apicalmente y como muchos 
otros miembros de las Echiteae, son conspicuamente rostradas, con el rostro 
delgado. 


MATERIALES Y METODOS 


Con el fin de determinar las posibles relaciones intergénericas de Bahiella, un 
total de 44 especies pertenecientes a 19 géneros de la tribu Echiteae (sensu En- 
dress & Bruyns 2000) con distribucion restringida al Nuevo Mundo, fueron 
incluidos en un analisis cladistico. Como grupos externos fueron escogidos dos 
representantes de las tribu Apocyneae (Beaumontia, Odontadenia) y seis de la 
Mesechiteae (Allomarkgrafia, Tintinnabularia, Mandevilla), basado en estudios 
moleculares previos (e.g, Sennblad et al. 1998; Sennblad & Bremen 2002) que 
han sugerido las relaciones cercanas de Echiteae con esas tribus. Finalmente se 
utilizaron dos representantes de las Malouetieae (Pachypodium) con el objeto 
de enraizar el arbol. 

Cincuenta y seis caracteres representado ciento treint ay unestados fueron 
evaluados para cada uno de las especies Gaile i cae caracteres fueron 
seleccionados de estudios previos (Endress et al. 1996; Simoes et al. 2004; Will- 
iams 2002, 2004), mientras que otros son incluidos por primera vez. Los estados 
de caracteres fueron evaluados en especimenes de herbario, flores preservadas 
y material fresco cuando estuviera disponible, los que a su vez fueron 
corroborados en forma paralela con descripciones de literatura (Morales 1997 
a, 1997 c, 1997 d, 1999 a, 2002 a, 2002 b, 2005 a, 2005 b; Morales & Williams 
2004, Williams 1998, 2002; Woodson 1933, 1936). La explicacion de los caracteres 
que no hayan sido utilizados anteriormente en analisis cladisticos previos, asi 
como algunas discrepancias encontradas entre lo citado en la literatura y lo 
presente en los especimenes son brindados en el apéndice 1. 

Los caracteres morfologicos evaluados fueron organizados en una matriz 
(Tabla 1) utilizando Winclada, version 1.00.08 (Nixon 2002). Todos los caracteres 
fueron tratados con un mismo peso y en forma desordenada. Especies con 
caracteres multiestados fueron reconocidos como polimorficos. El andlisis 
heuristico incluy6 una ronda de 500 replicaciones de secuencia y adicion 
aleatoria, con la opcién “Multi TBR + TBR” seleccionada y con un maximo de 
10000 arboles a mantener, salvando un arbol por réplica. El soporte relativo de 
los nudos del arbol de mayor consenso estricto fue obtenido mediante el 
procedimiento de “bootstrap” implementado en Winclada, con un numero to- 
tal de 1000 réplicas, 10 réplicas de busqueda aleatoria para secuencias 
adicionales, un arbol conservado en cada réplica y con la opcion “TBR” activada, 
salvando un maximo de 100 arboles en cada r€plica. 


] 


Taa.a 1.Matriz de datos morfoldgicos de 56 caracteres usados en el andlisis cladistico. 


Especie 


Pachypodium lamerei 
Pachypodium geayi 
Beaumontia grandiflora 
Odontadenia matogrossana 
Tintinnabularia gratissima 
Tintinnabularia mortonii 
Alllomarkgrafia brenesiana 
Allomarkgrafia plumeriifolia 
Mandevilla hirsuta 
Mandevilla sagittarii 
Allotoonia turbinata 


Asketanthera dolichopetala 
Asketanthera ekmanniana 
Bahiella blanchetii 

Bahiella infundibuliflora 


Laubertia contorta 


Estados de caracter 


BEE 


wn 
wn 


Sa cep TS LS ee es ee es eS SS > > se 


(L)@2 WaIs/9¥O' LINE 


TaBLa 1.continued 


Especie 


Macropharynx renteriae 
Macropharynx spectabilis 
Neobracea bahamensis 


uteum 
Prestonia falcatosepala 
Prestonia quinquangularis 
Prestonia ipomaeifolia 
Prestonia longifolia 
Prestonia vana 
Rhabdadenia biflora 
Rhabdadenia macrostoma 


Thoreauea paneroi 


Estados de caracter 


01010 
01010 


25 30 
00001 42110 
001 42110 
00001 42110 
00001 12110 
00001 32110 
00001 42110 
00001 32110 
00001 02110 
00001 02110 
01111 42110 
01101 42110 
0111] 42110 
01111 42110 


00001 b2110 
00001 12110 
00001 12110 


Caracteres polimorficos representados por letras de acuerdo a la siguiente disposicién: 
/3 


a=0/1,b=1 
? = datos desconocidos. 


Vilainvad Sd TVaOW 


6£€ 


340 BRIT.ORG/SIDA 22 


=a 


1) 


RESULTADOS 


Un total de 45 arboles parsimoniosos de 184 pasos fueron obtenidos por el 
analisis heuristico, con un indice de consistencia de 0.4 y un indice de retencion 
de 0.75. El arbol de mayor consenso estricto, asi como los valores de “bootstrap” 
mayores 0 iguales a 50 son presentados en la Figura 1. En general, al igual que 
otros analisis cladisticos basados unicamente en datos morfoldgicos que han 
involucrado miembros de las Echiteae y Mesechiteae (e.g., Simoes et al. 2004; 
Williams 2004) el soporte de los clados basales es bajo y una gran politomia 
prevalece; solamente algunos clados y subclados terminales estan soportados 
con valores de “bootstrap” mayores a 50%. Por lo tanto, la discusi6n subsecuente 
se orientara a comentar la conformacion de los clados y subclados del arbol de 
mayor consenso estricto. 

En primera instancia es interesante la posicién de Neobracea sugerida en 
este cladograma. Este género ha sido tradicionalmente ubicado dentro de las 
Echiteae, pero aca aparece ubicado en un subclado relacionado con el grupo 
externo y no con el interno (las Echiteae). Aunque el propésito de este analisis 
no es probar la monofilia de las Echiteae, la ubicacion de Neobracea sugiere la 
necesidad de evaluar la monofilia de la tribu tal y como se encuentra 
conformada en la actualidad (segtn Endress & Bruyns 2000). 

Dentro del grupo interno, el primer clado esta conformado por los géneros 
Stipecoma, Cycladenia, Rhabdadenia, Angadenia y Pentalinon, dentro del cual 
Cycladenia y Rhabdadenia conforman un subclado y Angadenia y Pentalinon 
otro subclado. Esta misma relacion entre Angadenia y Pentalinon, habia sido 
sugerida anteriormente por Williams (2004), quién obtuvo resultados similares 
en un analisis orientado a determinar la monofilia del género Echites. 

El siguiente clado esta conformado por 4 subclados que conforman una 
gran politomia sin resolver. Un total de 4 subclados son reconocidos: el subclado 
de Fernaldia, el subclado de Asketanthera, el subclado de Echites y el ultimo de 
ellos conformado por 10 géneros, que a su vez se encuentran agrupados en 
algunos subclados: Iemnadenia, Allotoonia, el subclado de Bahiella y Hylaea, 
el subclado de Macropharynx y Peltastes, el subclado de Thoreauea y Thenardia 
y el subclado de Prestonia y Laubertia. Sin embargo, a pesar de que los resultados 
sugieren que Bahiella esta relacionada con el resto de las Echiteae, basado en el 
bajo soporte de los clados basales y la gran politomia sin resolver prevaleciente, 
no es posible determinar por el momento y de manera concisa los géneros mas 
cercanamente relacionados con Bahiella. Partiendo de los resultados sugeridos 
por este analisis, es claro que son necesarios estudios moleculares 
complementarios para definir apropiadamente las relaciones intergénericas 
dentro de las Echiteae, asi como para probar la monofilia de la tribu. Analisis 
filogenéticos basados en caracteres morfoldgicos en otras tribus de las 
Apocynoideae (e.g., Mesechiteae), han resultado en cladogramas pobremente 
resueltos, en contraposicion con analisis basados en estudios moleculares (o 


MORALES, BAHIELLA E 


stipecoma peltigera 


mbellatus 
Echites yucatanensi 
Temnadenia stenantha 
a by oe ) i A, if, 


77 Allotoonia turbina 
Allotoonia noodsonan 
65 Bahiella blanchetii 
Bahiella ifnuitor 
83 lylaea a 
Hylaea le, ie 
Macropha ete 
38 Macropharynx Esuebie 
85 Peltastes p Itatu. 
a 


gf i 
S 


Thenardia galeottiana 
Pestone falcalosepala 


90 La nee boissien 


e ta 
ee ie so ifol 
ia Hise sk ue a ae 


Allomarkarai Bienesiane 


Tintinnabularia aan 
50 Tin tinnabulan ja mo. 
hireuta 


Mandevilla eagle 


SE geayi 
Pachypodium lamerei 


Fic. 7. Arbol d ‘oe er eae eee ad | AC deh nl 


y 4s parsimoni Valores de “bootstrap” mayores 
0 igual a 50 son indicados sobre las ramas. 


combinados), que han propuesto ee con mucho mas soporte (Simoes 
etal. 2004). En todo caso, los mismos estudios han conllevado 
a la necesidad de variar la configuracion original propuesta por Endress & 
Bruyns (2000) para conformar tribus naturales y monofiléticas. Sin embargo 
aun falta trabajo para determinar la conformacion definitiva de otras (e.g., 


342 BRIT.ORG/SIDA 22(1) 


Apocyneae), dado que se ha determinado que su constituci6n actual no es 
monofilética. 


TRATAMIENTO TAXONOMICO 

Bahiella J.F. Morales, gen. nov. Tipo: Bahiella blanchetii (A. DC.) JF Morales 
Fruticosa volubilis; folia opposita petiolata, coriacea, eglanduligera, revoluta; inflorescentia 
corymbosa, lateralis, pedunculata, multiflora; corolla regularis 5-partita, infundibuliformis vel 
hypocraterimorpha, corona et coronae lobis absens, antheris sagitattis, basi acuminata. Fructus 
folliculis divaricatus. 
Lianas con tallos viejos lenosos, con secrecion lechosa al cortarse, las ramitas 
ee porns teretes a subteretes, algo aplanadas en los nudos, mas rara- 

3 lé i iolares inc onspl¢ uos Hojas opuestas, peciola- 
das, sin soiree enel nervio central fae aieneate sin domacios abaxialmente. 
Inflorescencias cimosas, axilares, pedunculadas, con muchas flores, densamente 
aglomeradas distalmente, el raquis usualmente noduloso, bracteas florales 
escariosas, bracteolas ausentes; sépalos escariosos, con un coléter solitario, 
episépalo, dispuesto en la base de la cara adaxial, usualmente profunda e irregu- 
larmente lacerado, a veces casi totalmente dividido en varios segmentos; corola 
infundibuliforme o hipocrateriforma, glabra, glabrescente 0 inconspicuamente 


puberulenta exteriormente, sin estructuras coronales interiormente, el limbo 
actinomorfico, dext luto, estambres incluidos, las anteras con- 
niventes y aglutinadas en dos puntos a la cabeza estigmatica, las bases estériles 
conspic das y sagitadas, ovario bicarpelar, los carpelos unidos 
en forma apical al estilo, los 6vulos numerosos, la cabeza estigmatica fusiforme, 
con un anillo membranoso basalmente, nectario anular, irregularmente 
pentalobulado. Frutos apocarpicos, compuestos por dos foliculos, mas o menos 
divaricados, continuos, glabros, con diminutas costillas longitudinales; semillas 
numerosas, secas, rostradas, eee comosas apicalmente. 

Bahiella es un génerod especies, endémico a las formaciones costenas 
del estado del Bahia, Brasil. Su nombre genérico hace referencia al estado de 
Bahia, Brasil, la (nica localidad conocida por el momento. 


CLAVE PARA LAS ESPECIES DE BAHIELLA 


1. Corola hipocrateriforma, con la fauce de 2,5-3,5 mm de didmetro; ldbulos de 
la corola de 7-11 X 4-6 mm; anteras de 5,5-6 mm de largo; sépalos de 1-1,3 X 
J1-1,2 mm B. blanchetii a DC.) J.F. Morales 
. Corola infundibuliforme, con la fauce de 9-10 mm de didmetro;lébulos de 
de 14-16 X 10-13 mm, anteras de 77,5 mm de largo; sépalos de 2,2—3 
mm et ae J.F. Morales 
1. Bahiella blanchetii (A. DC.) J.F Morales, comb. nov. (Fig. 2). BAsIONIMo: Echites 
blanchetii A. DC,, Prodr. 8:448. 1844. Mitozus blanchetii (A. DC.) Miers, Apocyn. S. Amer. 219. 
1878. Tipo: BRASIL. BAHIA: Bahia, 1840 ({D, Blanchet 3223a (HOLOTIPO: G-DC, foto F neg. 7567, 
INB). 


as 


MORALES, BAHIELLA 343 
E 
> mm lcm ; f 
\ ene RY 
\\ Ih 5mm 
: iis Wi 
, = YY a 
Y VY 
Fic. 2 Bahiella blanchetii (A-G de Santos 2219, CEPEC; H—J de Amorin et al. 1386, CEPEC). A. Ramita con infl 
Natalla do lal pee dee pee ean [eee pranted} tnt 1.4 Ce | Lig fetal 4 Snalo.mostrando 
I colét lat D.Tubo del lal g linal biert t Jo | estilo y nectario. E Antera 
ista dorsal y lateral. F. Cab tica. G. Nectario. H. Foliculos. 1. Detalle de | Je los foli J. Semilla 


J 


344 BRIT.ORG/SIDA 22(1) 


Liana, los tallos viejos lenosos, ramitas usualmente teretes a subteretes, glabras 
a glabrescentes, coléteres interpeciolares ca. | mm de largo. Hojas: peciolos de 
6-13 mm de largo; laminas de 8-16,5 x 4,5-7,5 cm, coriaceas a subcoridceas, 
elipticas a obovadas u obovado-elipticas, redondeadas, obtusas a 
inconspicuamente retusas 0 apiculadas apicalmente, redondeadas, obtusas a 
raramente cuneadas basalmente, Seen conspicuamente revolutas 
marginalmente cuando viejas, esencial ,sin coléteres 
en el nervio central ad sialmente la venacion secundaria apenas evidente o 
inconspicua en ambas superficies, la venacion terciaria no evidente. 
Inflorescencias inconspicua y esparcidamente puberulentas a glabrescentes, 
pedinculo 4-65(-100) mm de largo, el raquis conspicuamente noduloso, 
pedicelos de 12-25 mm de largo, bracteas de 0,5-1 x 0,5 mm, escariosas, sépalos 
de 1-1,3 x 1-1,2 mm, levemente imbricados basalmente, angostamente ovados, 
agudos a obtusos apicalmente, muy esparcida e inconspicuamente puberulentos 
lé 0,5mm de largo, irregularmente 
lacerados corola hipocrateriforma, el tubo ptirpura intenso, rosado a rojizo, los 
lobulos lila a rojizos, pero blanquecinos marginalmente, glabra, glabrescente a 
inconspicua y esparcidamente puberulenta exteriormente, tubo de 15-17 mm de 
largo, de 2,5-3,5(-4) mm de diametro en el orificio, inflado sobre la posicion de 
los estambres, oe de 7-11 x 4 ° mm, angostamente obovados, extendidos 
ero vari t 
dorsalmente, cabeza estigmatica dé 15 L75 mm de largo; ovario de 1-2 mm de 
largo, glabro, nectario ca 1,6-2 mm de largo, anular, leve e irregularmente 


q glabrescen tesexteriormente 


-anteras de 5,5-6 mm de largo glabras 


pentalobula o0.F t por dos folic ulos apoca arpicos, de 14- 19x 0,5- 
0,7 cm, mas 0 menos divaricados, BIAPOS; continuos, con numerosas costillas 
longitudinales e i IncOnNnsf 1 uas; semi! llasdel5- 20mm de largo, coma apical de 2 8- 


4 cm de largo, color canela 

Distribucién, habitat y ecologia.—Endémica del estado de Bahia, Brasil, 
donde se encuentra en bosques humedos, vegetacion costera, vegetacion de 
restinga, y areas de vegetacion alterada asociada, en elevaciones de 0-100 m. 
Especimenes con flores han sido recolectados entre enero y febrero y de 
septiembre a noviembre. Material con frutos se reporta en enero, febrero, 
septiembre y noviembre. 

Bahiella blanchetii es una especie ayant comun, que se reconoce 
al instante por sus flores con corolas hipocrateriformas. Es interesante notar 
como este género y en particular esta especie, la cual fue descrita desde hace 
mas de 150 anos, ha permanecido desapercibida desde entonces. En forma gen- 
eral, pocos taxOnomos vieron el tipo, pues ni Miers (1878) ni Woodson (1936) lo 
examinaron y aparentemente solo Miller Argoviensis (1860) lo hizo, pero en 
todo caso lo traté en el entonces inclusivo género Echites. En todo caso, las 
colecciones disponibles antes de 1970 eran muy escasas y ayaa esta 
especie se conocia solo por el tipo y as por Regel 


MORALES, BAHIELLA E : 345 


y Riedel, ambas sin localidad ni fecha exacta y depositada en el herbario de la 
Universidad de Cambridge, Inglaterra (CGE) y en el Museo de Historia Natural 
de Viena (W). La semejanza de esta taxon con un Mesechites (i.e, Mesechites 
mansoanun (A. DC.) Woodson) es muy notable, ya que ambas especies son 
morfoldgicamente muy similares; inclusive, el especimen depositado en Viena 
fue examinado por Woodson y catalogado como M. mansoanus; sin embargo, 
como se explicé anteriormente, las caracteristicas de la cabeza estigmatica de 
ambas géneros son bien distintas y excluyentes, asi como otros caracteres 
morfologicos adicionales. En los ultimos 30 anos el incremento de la actividad 
botanica en Bahia, ha aumentado considerablemente el numero de especimenes 
disponibles hoy en dia, lo que ha permitido determinar en forma definitiva, el 
estatus ce de esta especie. 


BRASIL. Bahia: [|héus, Fazenda Jaguariba, al O de carretera Ilhéus-Olivenga, 
31 ie 1997 (fl), Arbo et al. 7835 (CEPEC, CTES), ie reserva biolégica do Mico-leao (IBAMA), entre 
Ilhéus y Una, 14 Sep 1993 (£1, fr), Amorin et al. 1386 (C C), Marat, carretera BR 030, entre Ubaitaba 
y Marat, 25 Feb 1980 (fl, fr), Carvalho et al. 166 ae. Cabralia, reserva biologica DO Pau-Brasil, 5 
Ene 1972 (£1, fr), ee 126 (CEPEC): S de Cumuruxatiba, ca Prado, 18 Ene 1977 Harl ey 
18089 (CEPE ce Z),e Una y Ilhéus, 23 Ene 1977 (£1), ee et a 18178 (CEPEC, INB, K, NY); 
Santa Cruz de ae carretera antigua, 21 Oct 1978 (f)), Mori et al. 10919 (CEPEC, a 
Nov 1971 (£1, Santos a2) (CEPEC); - Valenca, entre Valenca y Guaibim, 21 Feb 1975 (£1, fr), ae 
2888 (CEPEC): Itacaré, Nd le, caminoa Itacaré, 15 Nov 1992 (fl), Thomas et al. 9507 (CEPEC 
NY, USF). Datos perdidos: (f1), Regel s.n. (CGE); (fD, Riedel s.n. (U, W). 


72 pene pen dibabtloea Morales, sp. nov. (Fig. 3). Tipo: BRASIL. BAHIA: Ilhéus, 
nco da Vitoria (5.7 km al O del puente sobre el rio Fundao, camino a Ita en 
camino ee sector O de Mata da Esperanga, 15 Ene 1995 (fl), Thomas et al. 10765 ees 
CEPEC). 
A B. blanchetii (A. DC.) J.E Morales, cui affinis llae infundibulif is, lobis 14-16 mm longis, et 
antheris 7-7,5 mm longis differt. 


Liana, los tallos viejos lenosos, ramitas usualmente teretes a subteretes, glabras 
a glabrescentes, coléteres interpeciolares ca. 1 mm de largo. Hojas: peciolos de 
5-8 mm de largo; laminas de 8-16,5 x 3,8-5,5 cm, subcoriaceas, elipticas a 
obovado-elipticas, obtuso-apiculadas apicalmente, obtusas y cortamente 
atenuadas basalmente, revolutas marginalmente cuando viejas, glabras, 
eglandulares, sin coléteres en el nervio central adaxialmente, la venacion 
secundaria apenas evidente en ambas superficies, la venacion terciaria no 
evidente. Inflorescencias diminuta y esparcida a moderadamente papilado- 
puberulentas, pedtnculo de 95-96 mm de largo, el raquis inconspicuamente 
noduloso, pedicelos de 10-13 mm de largo, bracteas de 1-1,5 « 0,5-1 mm, 
escariosas, ae de 22. e x 1,9- e 6 mm, aha dunloyaler eles basalmente, 
angostamente ovad esparcida 
a moderadamente puberulentos exteriormente, mas raramente glabrescentes, 
coléteres ca. 1 mm de largo, irregular y profundamente lacerados; corola 
infundibuliforme, magenta basalmente, con los lébulos verde palido, glabra, 


346 BRIT.ORG/SIDA 22(1) 


Wi 2mm 
1G a 


a mm 


1.5 cm 


5cm 


Fic. 2. Rahiella infiundibylifl (Tt tal. 10765, CEPEC).A.R +. gd ia R Detalle delah davial 


dala lamina fali + dnl d lot C. Vista adaxial ded inal + dal lot | 


base. D. Detalle del tubo de | la. E. Antera, vista d ly lateral. F. Cab ti Atica. G. Ovario y nectario 
é 


MORALES, BAHIELLA, FE E 347 


glabrescente a inconspicua y esparcidamente puberulenta exteriormente, la 
parte inferior del tubo de 10-12 x 2,7-3,2 mm, recto, la parte superior de 10-12 
mm de largo, de 9-10 mm de diametro en la fauce, l6bulos de 14-16 x 10-13 
mm, obovados, extendidos pero variadamente reflexos marginalmente; anteras 
de 7-7,5 mm de largo, glabras dorsalmente, cabeza estigmatica de 2-2,2 mm de 
largo; ovario de 1,8-2,1 mm de largo, glabro, nectario de 1-1,3 mm de largo, anular, 
leve a moderadamente pentalobulado. Foliculos desconocidos. 

Distribucién, habitat y ecologia.—Bahiella infundibuliflora es conocida 
unicamente de la localidad tipo, donde crece en bosques hmedos y margenes 
de bosques, en elevaciones bajo 50 m. El unico especimen con flores conocido 
fue recolectado en enero. 

Bahiella infundibuliflora es morfologicamente muy similar a B. blanchetii 
y aunque de primera instancia, ambas especies pueden confundirse, se pueden 
diferenciar con facilidad. De esta manera, B. infundibuliflora se puede distinguir 
facilmente por su corola infundibuliforme (vs. hipocrateriforma), con la fauce 
del tubo de 9-10 mm de ancho (vs. 2,5-3,5(-4) mm), sépalos mas largos y anchos, 
anteras de 7-7,5 mm de largo (vs. 5,5-6 mm de largo) y lobulos de la corola mucho 
mas grandes y desarrollados, de 14-16 x 10-13 mm (vs. 7-11 x 4-6 mm). 


APENDICE 1 
CARACTERES Y ESTADOS DE CARACTER UTILIZADOS EN EL ANALISIS 
CLADISTICO, CON ALGUNAS NOTAS EXPLICATORIAS 


A continuacion se citan los caracteres y estados de caracter utilizados en el 
analisis cladistico. Notas explicatorias son incluidas para aquellos que sea 
necesario, asi como cuando se hayan encontrado discrepancias entre lo citado 
por la literatura y lo encontrado entre los especimenes. Los caracteres 1, 3, 7, 8, 
20, 23, 26, 38, 44, 45, 46, 47, 48, 49, fueron tratados por Simoes et al. (2004), 
mientras que el 12, 13, 14, 16, 28, 29, 36, 40, 50, 52, 53, 55 por Williams (2004). Por 
lo tanto, no seran explicados aqui. 


1. Habito 4. Tallos inferiores 
0 Liana o hierbas escandentes 0 Herbaceos, no lignificados 
1 Hierba perennes 1 Variadamente lignificados 
2 Arboles 0 arbustos Casi sin excepcidn, la mayoria de los 
2. Tallos con espinas miembros neotropicales de las Echiteae o 
0 Si Mesechiteae son lianas, con los tallos viejos 
variadamente lenosos, algunos veces con la 
En general los miembros de las Echiteae y corteza conspicuamente suberosa. Sin em- 
Mesechiteae no tienen espinas 0 aguijones bargo, Cycladenia es una hierba erecta o 
en el tallo, pero este en un cardcter que se suberecta, que se caracteriza por sus tallos 
presenta en forma ocasional en géneros de herbaceos, no lenosos (Woodson 1936). 
otrasa tribus (e.g., Malouetieae) 5. Yemas axilares cubiertas por grupos de 
3. Latex coléteres filiformes 
0 Lechoso 0No 


1 Acuoso 15S! 


En general, la pubescencia de los tallos y 
brotes jovenes de las Apocynaceae es muy 
variable, pero en las Echiteae Unicamente en 
Stipecoma se da la presencia de coléteres 


tiliformes envolviendo las yemas axilares 


1 Opuestas 
7. Hojas con coléteres en nervio central 
0No 


1Si 
8. Hojas con domacios 
No 


1Si 
9. Hojas peltadas 
0 Ausentes 
1 Presentes 
10. Consistencia de las hojas 
0 Coriaceas 
1 Membrandceas 
11.Margenes revolutos 
0 Ausentes 
1 Presentes 
12. Venas secundarias impresas 
0Si 


1No 
13. Venacion terciaria en la cara abaxial 
0 Conspicua, bien impresa 


1 Inconspicua, apenas impresa O no impresa 
14. Venas terciarias finamente reticuladas 
0s 


f 
1No 


15.Venas terciarias perpendiculares al 
| 


nervio centra 
Si 
1No 


Las venas terciarias Cuando presentes, tienen 


patrones irregulares en cuanto 


goon ag. en ‘aia ee (e.g., 


BRIT.ORG/SIDA 22(1) 


En forma general, de igual manera que se 
presenta en otras tribus relacionadas (e.g,, 
Mesechiteae), los miembros de las Echiteae 
tienen una sola inflorescencia por nudo, 
pero en unos pocos géneros (e.g., Macro- 
pharynx, Peltastes), dos inflorescencias por 
nudo usualmente estan presentes 


18. Tipo de inflorescencia 


acimo 
1 Cima sin ramificar o con una sola division 
basal 
2 Cima ramificada 
Williams (2004) menciond que las 
inflorescencias en Laubertia contorta (M. 
Martens & oe ede no son 
ramificadas, per fl encias 
con al menos una ramificacion también son 
encontradas en esta especie, por lo que este 
estado de caracter debe tratarse como 
Eon eNice. 


19. Fl g I pect beli 


forme 
0Si 
1No 


20. Bracteas 


0 Escariosas, inconspicuas 


1 Foliaceas, conspicuas 


21. Bracteolas 


1 Presentes 

La presencia de bracteolas es raro en las 

Echiteae y Mesechiteae, presentandose 

Shes en Macropharynx y Peltastes, 
nque si bien, bracteolas inconspicu 

ie presentarse en unas pocas especies 

de Mandevilla o Neobracea 


22. Consistencia de los sépalos 


Hlalleida 


1 
mas © menos perpendicular en relacion al 


nervio centr 
16. Posicion de : inflorescencia 

0 Axilar 

1 Terminal 
17. Numero de inflorescencias por nudo 


na sola inflorescencia en alguna de las 


dos axilas 
1 Dos inflorescencias, una en cada axila 


0 Escariosos 
1 Foliaceos o subfoliaceos 


23. Disposicioén de los coléteres 


0 Solitarios y epicentricos 
1 Filas continuas e ea tee en la base 
2 Alternos con los sépalos 
Woodson (1936), en su monografia de 
Pentalinon (tratado entonces como 


p= ae 


Ure hi tes 5) me nci ion ie, que e los léteres en P. 
andrieuxti (Mall. Arg.) B. Hansen & Wunderlin 
eran alternos con los sépalos, pero la 
evaluacion de dicho caracter en diferentes 


MORALES, BAHIELLA 


especimenes han revelado que estos se 
encuentran distribuidos de manera con- 
tinua en una hilera de forma irregular a lo 
largo de la base del sépalo. 
24. Numero de coléteres por caliz 

0 Cero 


1 Cinco 
2 Seis O mas 


25. Forma de la corola 


0 Hipocrateriforme 
1 Infundibuliforme, parte superior conica o 
tubular 


re q | 


3 Urceolada 

4 Rotada 

Tanto Simdes et al. (2004) como Williams 
(2004) han propuesto estados de caracter 
que difieren en la clasificacién de corolas 
infundibuliformes, siendo homédlogos en lo 
referente a corolas hipocrateriformes o 
rotadas. En este andlisis, las oe 
infundibuliformes fueron divididas S 
estados,el primero para clasificar las ee 
con la parte superior del tubo cédnica y el 
segundo para aquellas con la supe- 
rior campanulada. Dentro de las Echiteae, 


7 t ~ ' 
con facilidad por la parte superior de la 
corola (e.g.,Angadenia, Pentalinon),la cual es 
siempre campanulada 


26. Parte inferior del tubo 


0 Recto 
1 Curvado 
27.Boton floral con union Id6bulo-tubo 


1 Presente 
La presencia de coronas anulares, limitadas 


a unos pocos géneros en las Echiteae (Mo- 
rales 1997 a, b, 2002 a; Williams 2002; 
woodsell2s ) provecan ginacion 


| | if 
UNO 


con el tubo en gorones iietales que se 
presenta de Laubertia 
y Prestonia, pero que no es perceptible en 
Thoreauea (Morales 2005 b; Williams 2002). 


28. Corona anular 


0 Ausente 
1 Presente 


29. Corona supraestaminal 


0 Ausente 

1 Presente 

En ie Echiteae. a colon epiestaminal 
J 


C k F por cinco 
Sn dj 4 folia ubicados 


en forma opuesta a cada antera 


30.Margen de los Iébulos en botones 


florales 
0 Enroscados en espiral sobre su eje varias 
ces 


1 Enroscados en espiral una sola vez 
Los Idbulos de la corola en las Echiteae y 
Mesechiteae, se encuentran enroscados en 
espiral, pero de manera regular estan 
invaginacos hacia el interior,de manera que 
el margen externo de los ldbulos se ven 
sll cba en ik ee externa Uni- 
traposicion en 
Asetonthera, los guile estan enroscados 
sobre su eje hacia el apice del botédn varias 
ee manera que el margen externo de 
lo ldbulos se ven enroscados varias veces 
sobre su eje 


31. Color predominante de los Idbulos de la 
orola 


0 Amarillo 
1 Rosado 
2 Rojo o morado 
3 Verde 
4 Blanco o crema 


32. Forma de los I6bulos 


2 Obovados 

En forma general, la mayoria de las géneros 

y especies de las Mesechiteae y Echiteae 

tienen Idbulos oblicuamente obovados, 
rf + | + 7 lAkK | A 


ofiliformes es caracteristico para unos pocos 
Woodson 1936; Williams 2002; Morales & 
Williams 2004 


— 


33.L6bulos erectos o suberectos 
0 Si 


1 No 

Los Idbulos de la corola se encuentran 
usualmente extendidos de manera irregular 
en la mayoria de los géneros de las Echiteae 
y Mesechiteae. Sin embargo, lobulos estricta- 


34. 


36. 


37. 


38. 


39. 


mente erectos o suberectos, estan limitados 
a unas pocas especies de Mandevilla y al 
género Thoreauea (Williams 2002; Morales 


2005, b). 
Lobulos torcidos en espiral distalmente 
Si 


1No 

Los oe en ie eee eae son 

antésis, pero 
Ponelnnneets reflexos de manera irregular 
en sus bordes; los Idbulos erectos o 
suberectos son menos comunes. Por otro 
lado, la presencia de lObulos enroscados en 
espiral distalmente solo esta presente en 
Allotoonia (Morales & Williams 2004 

ee fe city gaa 


0 Ausente o inconspicua, los pelos hasta 1 
mm de largo 
1 Presente y conspicua, los pelos mas de 2 
e largo 
Aunque la pubescencia de la corola como 
tal puede ser variable en las Echiteae, 
unicamente en Fernaldia se da la presencia 
de pelos largas y blancos en la cara adaxial 
de los ldbulos (Morales 2002 a). 
Tubo de la corola 
0 Diminuto, menos de 6 mm de largo 
1 Pequeno, 8-19 mm de largo 
2 Mediano, 20-33 mm de largo 
3 Grande, mas de 35 mm de largo 
Insercion de los estambres 
0 Mitad distal, cerca de la boca 
1 Mitad basal, cerca de la base 
Filamentos 
nteras + sésiles 
1 Menos de 1 cm de largo 
2 Mas de 3 cm de largo 
Filamentos envolviendo totalmente al 
fo) 


0 Si 

1No 

Los filamentos de la mayoria de las Echiteae 
y otras tribus de las Apocynoideae son muy 
cortos y casi ausentes, por lo que apenas 
tienen contacto con el estilo en la regién 
distal de este, ue antes de la posicioén de 
la cabeza estig 
generos (e.g., Forsteronia, Thenurdia: 
Thoreauea), dado el pequeno tamano del 


Cin embarg enotro 


BRIT.ORG/SIDA 22(1) 


tubo y estilo envuelven a este ultimo casi 
en la totalidad de su longitud. 


40. Anteras 


41. 


42. 


44. Canales 


0 Incluidas 
1 Apicalmente exsertas 

asi totalmente exsertas 
Pubescencia del apice de las anteras 
0 Ausente 
1 Presente 
La pubescencia dorsal de las anteras ha sido 
utilizado en otros andlisis (e.g., Simdes et al. 
2004), pero este cardcter puede variar 
hotablementes en as eae Sin em- 


bargo, | 

restringida a la region estéril y apical de las 
anteras, que se puede incluso prolongar en 

un apéndice. 

Apice de las anteras 

0 Triangular,no prolongado en un apéndice 
filiforme 

1 Prolongado en un apéndice filiforme 

En general, la mayoria de las Apocynoideae 

neotropicales tienen los apices de las 

anteras variadamente triangulares, ya sea 

agudos 0 acuminados. Sin embargo, la 

prolongacion de estos apéndices por varios 

milimetros es raro y esta limitado a unos 

pocos geéneros 


43.Apéndices filiformes de las anteras 


ier en espiral 

0 Aus 

1 ee 

Los ee filiformes de las anteras 
ando | pueden ser erectos (€.g., 

= er e) de, entrelazados en 

espiral (Pentalinon). 

guia de las anteras 

0 Compuesto principalmente de engrosa- 

mientos endoteciales 
1 Compuesto principalmente de esclerén- 


present 


quima 
45. Base de las anteras 


0 Acuminadas 
1 Agudas 
2 Obtusas a redonde 


adas 
46. Union antera/cabeza estigmatica 


0 Anteras unidas por un parche circular de 
células con apariencia de tricomas 

1 Anteras unidas por un aro de pelos en 
forma de herradura 


MORALES, BAHIELLA f 7 TeTARANED es 


2 Anteras unidas por un aro de pelos en 53. Grosor de los foliculos 


forma de herradura y una tira longitudi- 0 Gruesos, mas de 10 mm de ancho 
nal delgada 1 Delgados, hasta 9 mm de ancho 
3 Anteras unidas por fusién celular 54. Folicul I tillas longitudinal 
47. Cabeza estigmatica en corte transversal 0 Si 
0 Circular o subcircular 1N 
1 Pentagonal 55. Semillas 
2 Con cinco costillas prominentes 0 Rostradas 
48. Costillas de la cabeza estigmatica 1 Truncadas 
0 Restringidas a la base 56. Extremo micropilar de la semilla 
1 Distribuidas a lo largo de la cabeza estig- 0 Extendida mds alla del punto de naci- 
matica miento de la coma 
49. Anillo en la base de cabeza estigmatica 1 No extendida mas alla del punto de 
0 Ausente nacimiento de la coma 
1 Presente La presencia de una coma micropilar en la 
50. Nectario semilla es algo comun en muchas Apocy- 
0 Anular noideae.Generalmente, el extremo micropilar 
1 Dividido en cinco glandulas termina en el punto en el cual la coma nace. 
51.Gineceo En muy pocas especies, el extremo 
0 Apocarpico micropilar se prolonga por varios mm por 
1 Sincarpico encima del punto de insercién de la coma, 
52. Folicul convirtiéndose en un caracter potencial 
0 Moniliformes o torulosos para la delimitacién genérica e incluso tribal. 
1 Cilindricos 


AGRADECIMIENTOS 


Un reconocimiento y agradecimiento profundo al Herbario del Centro de 
Pesquisas do Cacau (CEPEC) en Bahia, Brasil, por el numeroso envio de mate- 
rial en préstamo, que permitio definir y clarificar el status definitivo de este 
nuevo género, lo que de otra manera no hubiera sido posible. Por otro lado los 
siguientes herbarios colaboraron con el préstamo y uso de colecciones: CGE, G- 
DC, K, NY, P, W. Un agradecimiento sincero a Fabian Michelangeli (NY) por su 
paciencia y disposicion a clarificar varias dudas del andlisis cladistico aca 
presentado. También reconozco la revision critica y numerosas sugerencias de 
Mary Endress (Z) en muchos aspectos de este trabajo, lo que permitio su 
correccion y mejoria notablemente. También se agradece la ayuda de Lucille 
Allorge por facilitar el acceso y estudio del material en los herbarios del Museo 
de Historia Natural de Paris, Francia, asi como de Bruno Wallnéfer por las 
mismas razones en el Museo de Historia Natural de Viena (W), Austria. 
Finalmente, quiero agradecer a Cecilia Ezcurra (BCRU) ya un revisor anonimo 
por la revision critica del manuscrito. 


REFERENCIAS 


Brown, R. 1810. On the Asclepiadeae, a natural order of plants separated from the 
Apocineae of Jussieu. Mem. Wern. Nat. Hist. Soc. 1:12-78. 


352 BRIT.ORG/SIDA 22(1) 


Canoolte, A d&. 1844. Apocynaceae. In: A de Candolle, ed. Prodromus systematis naturalis 
regni vegetabilis, Vol. 8. Treuttel & Wurtz, Paris. Paris. Pp. 317-489. 

Enoress, M.E., B. Sennetao, S. Nicsson, L. Civeret, M. CHASE, S. Huysmans, E. Grarstrom & B. BREMER. 
1996. A phylogenetic analysis of Apocynaceae s. str. And some related taxa in 
Gentianales:a multidisciplinary approach. Opera Bot. Belg. 7:59-102. 

Enoress, M.E. & P. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 
66:1—56. 

Hansen, B. 1985.A monographic revision of Forsteronia (Apocynaceae). Ph.D. dissertation, 
Univ. of South Florida, Tampa. [University Microfilms International, Ann. Arbor] 

Jupp, W., W. Sanoers & M. Donochue. 1994. Angiosperm family pairs: preliminary phyloge- 
netic analyses. Harvard. Pap. Bot. 5:1-51 

Miers, J. 1878. On the Apocynaceae of South America. Williams & Norgate, London, 
Edinburgh. 

Mora es, J.F.1997a.A synopsis of the genus Prestonia (Apocynaceae) seccion Jomentosae 
in Mesoamerica. Novon 7:59-66. 

Morates, J.F.1997b.A reevaluation of Echites and Prestonia section Coalitae (Apocynaceae). 
Brittonia 49:328-336. 

Morates, J.F. 1997c. A synopsis of the genus Allomarkgrafia (Apocynaceae). Brittonia 49: 
337-345 

Morates, J.F.1997d [1998].A synopsis of the genus Macropharynx (Apocynaceae).Rhodora 
99:58-68. 

Morates, J.F. 1999a. Hylaea (Apocynaceae-Apocynoideae), a new genus from South 
America. Novon 9:83-85. 

Morates, J.F. 1999b. A synopsis of the genus Odontadenia (Apocynaceae), In AJ.M. 
Leeuwenberg, ed. Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67: 
381-477. 

Morales, J.F. 2002a. Studies in Neotropical Apocynaceae |:A revision of the genus Laubertia 
Rhodora 104:170-186. 

Monrates, J.F.2002b. Studies in Neotropical Apocynaceae ll:A revision of the genus Fernaldia. 
Rhodora 104:186-200. 

Morates, J.F. 2003. Studies in Neotropical Apocynaceae Ill: A revision of the genus 
Secondatia, with discussion of generic classification. Candollea 58:305-319. 

Morates, J.F.2004. Estudios en las Apocynaceae Neotropicales VIl: Novedades taxonomicas 
en Prestonia (Apocynaceae, Apocynoideae) para Colombia y Ecuador,con comentarios 
sobre el grado de lobulacién del nectario. Candollea 59:159-165. 

Monrates, J.F. 2005a. Estudios en las Apocynaceae Neotropicales XIll: revisién del género 
Temnadenia (Apocynoideae, Echiteae). Candollea 60:207-231. 

Morates, J.F. 2005b. Estudios en las Apocynaceae Neotropicales XV: Sinopsis del género 
Thoreauea (Apocynoideae, Echiteae), con una nueva especie de Veracruz, México. 
Brittonia 57:258-263. 

Morates, J.F. & J. Wittiams. 2004. Allotoonia,a new neotropical genus of Apocynaceae based 
on a subgeneric segregate of Echites. Sida 21:133-158. 


MORALES, BAHIELLA E E Al 353 


Mutter Arcoviensis, J. 1860. Apocynaceae. En: C.F.P. von Martius, ed. Flora Brasiliensis 6(1). 
Munchen, Wien, Leipzig. Pp. 1-180. 

Nixon, K. 2002. Winclada, version 1.00.08. Programa de computacidn distribuido por el 
autor en, http://www.cladistics.com. Cornell University, Ithaca, NY. 

Poraieter, K & V. Avsert. 2001.Phylogenetic relationships within Apocynaceae s.|. based on 
trmL intron and trnL-F spacer sequences and propagule characters. Ann. Missouri Bot. 
Gard. 88:523-549, 

SENNBLAD, B. & B. Bremer. 1996. The familial and subfamilial relationships of Apocynacecae 
and Asclepiadaceae evaluated with rocL data. Pl. Syst. Evol. 202:153-175. 

SENNBLAD, B. & B. Bremer. 2002. Classification of Apocynaceae s.|. according to a new ap- 
proach combining Linnaean and phylogenetic taxonomy. Syst. Biol. 51:389-409. 

SENNBLAD, B., M.E. Enpress & B. Bremer. 1998.Morphology and molecular data in phylogenetic 
fraternity: the tribe Wrightieae (Apocynaceae) revisited. Amer. J. Bot.85:1143-1158. 

Simoes, A., M.E. Enpress, T. vAN DeR Niet, L. KinosHita & E. Conti. 2004. Tribal and intergeneric 
relationships of Mesechiteae (Apocynoideae, Apocynaceae): evidence from three 
noncoding plastid DNA regions and morphology. Amer. J. Bot. 91:1409-1418. 

Wiuams, J.K. 1998. A revision of Thenardia (Apocynaceae, Apocynoideae). Lundellia 1:7 


8 

Wiuams, J.K. 1999. A phylogenetic and taxonomic study of the Apocynaceae subfamily 
Apocynoideae of Mexico with a synopsis of subfamily Plumerioideae. Ph.D. disserta- 
tion. Austin: The University of Texas. 

Wituams, J.K. 2002. Thoreauea (Apocynaceae: Apocynoideae), a new genus from Oaxaca, 
Mexico. Lundellia 5:47-48. 

WILLIAMS, J.K. 2004. Polyphyly of the genus Echites (Apocynaceae: Apocynoideae:Echiteae): 
evidence based on a morphological cladistic analysis. Sida 21:117-132. 

Woopson, R.E. 1933. Studies in the Apocynaceae IV. The Américan genera of Echitoideae. 
Ann. Missouri Bot. Gard. 20:605-790. 

Woopson, R.E. 1935. Studies in the Apocynaceae. IlV.The American genera of Echitoideae. 
Ann. Missouri Bot. Gard.22:153-306. 

Woopson, R.E. 1936. Studies in the Apocynaceae. IV. The American genera of Echitoideae. 
Ann. Missouri Bot. Gard. 23:169-438. 


354 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


SusAN McCiure.1996. The Herb Gardener: A Guide for All Seasons. (Paperback 
edition). (ISBN 0-88266-873-0, pbk.). Storey Publishing, 210 MASS MoCA 
Way, North Adams, MA, 01247. U.S.A. pOnlets: # ooer 1-800-441-5700; 
wwwstorey.com). $14.95, 236 pp., color photogray d drawings, appen- 
dices, index, 8 3/8" x 10". 


Gardeners and cooks alike will learn a a deal from The Herb gardener: A Guide for All Seasons. 


Author Susan McClure has created a comprehensive compendium for people interested in herb g 
dening; subjects include a eae on designing, garden duties throughout the year, crafts ee 
projects, as well asa growing gui 

The book is divided into five main parts concerning a such as herbs, herb gardening 
Basics, seasonal planning in the garden, herb projects, and a grower’s guide. 
gardening basics chapter shows the reader items to pete in planning a garden and it 
also ie) in oe a garden space. General preplanning considerations are discussed including 


a 


plant life cycles, and plant tolerances. Also included are environmental considerations before you 
plant (sun, soil, fertilizer, pH, water, etc.). The portion of this chapter on designing your garden will 
be incredibly helpful information for most readers. The author has included many different garden 
types/shapes such as hardscape considerations, included labeling suggestions and maintenance 


thoughts. 

The section about yearlong gardening divides the year up into the four seasons and provides 
gardeners with typical, well-organized inf h snot occu: quae rach period of 
the year. One of the very helpful f in this bool he side | PI through- 


out the text; examples are the seed sprouting facts chart in the Spring period, harvesting techniques 
and troubling pests and diseases charts in the summer period, first frost estimator in the fall period, 
and plants for an indoor garden 1 in ah winter pene od. 

The section on recip d d | cool benefits of the pla 
ning and herbal harvest. Tie portion iaeludes auultiple methods of preserving herbs for longer-term 
hj 


use, good h tions, recipes, herbal vinegars, cheese and butters, a wreaths. 
It also say oie a helpful mre of nutritional value for the common her 
nal section of the book includes a very useful grower's guide. Ne section vee read- 


] 


ers with oe a along nh nace of 52+ eke al f 


J 


varieties as files for herbs are organized by Iso incl 
] 


opagation, cultivars, potential problems, harvesting, use, and more. Each profile 


ae summaries of oo in the forms of colorful icons for quick reference. 

If you have a hankering to make your own pesto or create your own herbal seasoning blend, 
check out The Herb Gardener: A Guide for Ail Seasons and plan your own herbal garden. McClure 
has written a text that is easy to read and easy to use. It’s full of helpful side boxes, charts and herbal 
profiles. This book would be a ae aeditiont to the pEary of gardeners, cooks and herbal connois- 
seurs.—Lee Luckeydoo, Herbariu h Institute of Texas, 509 Pecan Street, Fort Worth, 
TX 76102-4060, U.S.A. 


SIDA 22(1): 354. 2006 


ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XxV: 
NOVEDADES Y NUEVOS REPORTES EN LAS APOCYNACEAE 
(APOCYNOIDEAE: RAUVOLFIOIDEAE) DE VENEZUELA 


J.Francisco Morales 


Instituto Nacional de Biodiversidad 
Apartado 22-3100 
Santo Domingo, Heredia, COSTA RICA 


RESUMEN 


Se proponen varias Seine poniendiaturales en mere género Mandevilla, incluyendo una nueva 
especie (M. tristis J.F Morale anu dn (M. angustata (Steyermark) J.F Morales), y un 


nuevo sindnimo (M. bella a Woodson). Nuevas adiciones a la Flora de Venezuela en los géneros 
Aspidosperma (A. rigidum Rugby, A. schultesii Woodson), Mandevilla (M. vanheurckii (Mull. Arg.) 
Markgr.) y Odontadenia (O. killipii Woodson) son reportadas. 


ABSTRACT 


Several nomenclatural novellies are (piepered in the eeenus Mangers: la, including a new species (M. 
tristis J. Morales), a new k) J.-F. Morales) and a new synonym 
(M. bella (Pittier) Woodson). New additions to ache Venezuelan Flora in the genera Aspidosperma (A. 
rigidum Rugby, A. Sauultesty Wesdsen)) Mandeville AM NGDNEM TCE an Arg.) Markgr.), and 
OnE. clip lin the 
genus Mandevilla, incl species (M. tristis J.E Morales), a new combination (M. angustata 
Geom J.E Mo ouik a new report ((M. vanheurckii (Mull. Arg.) Markgr) and a new synonym 
(M. bella (Pittier) Woodson 


Venezuela es uno de los paises con una alta diversidad de Apocynaceae s. s.,con 
un estimado de 30 géneros y ca. 200 especies (Morillo 1989a). El numero de 
géneros puede variar dependiendo de la aceptacion de Iabernaemontana como 
una sola unidad genérica segtin lo propuesto por Leeuwenberg (1994) 0 dividido 
en una serie de grupos como lo propuesto por Allorge (1985). Asimismo, la 
relativa confusa taxonomia prevaleciente en algunos géneros como Aspi- 
dosperma y la divergencia de taxones aceptados de ese género en diferentes 
tratamientos floristicos puede incrementar o decrecer el numero estimado de 
taxones. 

En la ultima Buonep ae del género Mandevilla, Woodson (1933) report6 
un total de 15 esy [M. congesta (Kunth) Woodson, M. bella (Pittier) Woodson, 
M. caurensis Marker, M. fendl eri (Mull. Arg.) Woodson, M. hirsuta (Rich) K. 
Schum., M. javitensis (Kunth) K. Schum., M. lancifolia Woodson, M. moritziana 
(Mull. Arg.) Donn. Sm., M. scabra (Hoffmanns. ex Roem. & Schult.) K. Schum., 
M. subcarnosa (Benth.) Woodson, M. subsagittata (Ruiz & Pav.) Woodson, M. 
subspicata (Vahl) Markgr, M. surinamensis (Pulle) Woodson, M. veraguasensis 


SIDA 22(1): 355 — 365. 2006 


356 BRIT.ORG/SIDA 22(1) 


(Seem.) Hemsl., M. villosa (Miers) Woodson)| para Venezuela, de las cuales cuatro 
(M. bella, M. caurensis, M. lancifolia, M. moritziana) fueron consideradas 
endémicas. En forma posterior, mas especies f descritas (e.g., Markgraf 1971, 
1978; Monachino 1958, 1961; Woodson 1948) aumentando a 23 el namero 
aproximado de taxones conocidos para el pais. En su tratamiento de los géneros 
de Apocynaceae de Venezuela, Clausnitzer (1968) reporto un total de 22 especies, 
mientras que Morillo (1989 b), en conjuncion con la descripcion de dos nuevos 
taxones, report6 la presencia de M. annularifolia Woodson (anteriormente 
conocida solo de Colombia) y propuso dos nuevos sindénimos. En el tratamiento 
de las Apocynaceae para la Flora en la Guyana Venezolana, Morillo (1995) trato 
un total de 24 taxones, estimando un total de 40 especies para Venezuela. 
Aunque si bien este ultimo trabajo se trata de un tratamiento de flora para un 
area geografica especifica (y no para toda Venezuela), tiene una relativa 
importancia, por el hecho de incluir algunas especies previamente no conocidas 
en el pais. El tratamiento preliminar de una nueva monografia de Mandevilla 
(Morales, datos sin publ.) contabiliza un total de al menos 37 taxones en Ven- 
ezuela, algunos recientemente descritos (Morales 2005). De ellos, tan solo cinco 
pertenecen al subgénero Mandevilla, mientras que el resto pertenecen al 
subgénero Exothostemon (Woodson 1933). Como resultado, varias novedades 
son propuestas, incluyendo nueva sinonimia, un nuevo reporte, una nueva 
combinacion, y una nueva especie. 

Por otro lado, hasta el momento, no existe un tratamiento completo de la 
familia (s.s.) para el pais y Unicamente tratamientos para los géneros 
(Clausnitzer 1968) 0 regiones geograficas determinadas (e.g., Morillo 1989a; 
Zarucchi et al. 1995) han sido publicados. 

Mientras revisaba material sin identificar de Venezuela, varios taxones 

ente no reportados para ese pais de los géneros Aspidosperma 
(papi ies yO ren (Apocynoideae) fueron identificados, asi como 
una serie de novedades taxonomicas en Mandevilla (Apocynoideae) estas 
altimas derivadas principalmente de la elaboracién de una nueva monografia 
para el género. Estas novedades son presentados a continuacion, agrupados en 
subfamilias y tribus de acuerdoa la clasificacion propuesta por Endress y Bruyns 


Mandevilla angustata (Steyerm.) J.-F Morales, comb. & stat. nov. (Fig. 1). BAsionimo: 
Mandevilla subcarnosa var. angustata Steyerm., Acta Bot. Venez. 2:318. 1967. Tipo: VENEZU- 


ELA. BOLIVAR: Auyan-tepui, cumbre de la parte Norte, entre campamento Llovizna y pie del 
segundo Muro, 1] May 1965 ({]), Stevermark 93737 (HOLOTIPO: VEN; ISOTIPOs: MO (fotocopia en 

J, US, VEN). 
Lianas o arbustos erectos; ramitas jovenes levemente anguladas o aladas, 
subteretes con la edad, tallos sdlidos a ligeramente huecos, glabros, coléteres 
interpeciolares inconspicuos, hasta 0,8 mm de largo. Hojas opuestas; peciolos 


MORALES : 357 


Zi: 


De 


Fic. 1. Mandevilla angustata (Liesner & Holst 20762, VEN). A. Ramita fértil con flores y frutos. B. Detalle de la superficie 
Jo vtal da la hn} Pisani ea lot In] dol : tral. C. C4liz Jicelo v bra D. Vista adaxial de 


un sépalo, t Jo el col 
ici 


y teg t F. Antera, vista dorsal. G. Cat tig Stica.H Nectario y ovario. | Semilla 
J. Detalle de la pubescencia de las semilla. 


358 BRIT.ORG/SIDA 22 


1) 


3-8 mm de largo; laminas foliares 2,6-10 x (1-)1,4-3,9(-5) cm, obovadas a 
obovado-elipticas, el Apice obtuso, obtuso-apiculado a agudo, la base cuneada, 
los coléteres distribuidos en forma irregular a lo largo del nervio central, 
firmemente membranaceas, glabras o glabrescentes, usualmente no revolutas, 
venas secundarias y terciarias apenas impresas en ambas superticies. 
Inflorescencia ligeramente mas larga que las hojas subyacentes, subterminal, 
pero rapidamente volviéndose axilar, diminuta e inconspicuamente 
puberulenta cuando joven, pero volviéndose rapidamente glabra o glabrescente, 
con 5a 15 flores, sésiles o subsésiles, el pedtnculo (cuando raramente presente) 
de 1-5 mm de largo, pedicelos 3-5 mm de largo (a veces hasta 8 mm en 
fructificacion), bracteas 1-2 x 0,5-1 mm, angostamente ovadas, escariosas; 
sépalos 2-3 x 1,4-1,7 mm, ovados, obtusos a agudos apicalmente, el apice no 
reflexo, escariosos, glabrescentes externamente, diminuta e inconspicuamente 
ciliolados marginalmente, el coleter solitario, irregularmente lacerado 
apicalmente; corola infundibuliforme, amarilla, inconspicuamente papilada a 
glabrescente externamente, el tubo giboso basalmente, la parte inferior 27-32 
x 2-2,5 mm, la parte superior 13-19 mm de largo, conica, 14-22(-25) mm de 
diametro en la fauce, el apice del boton floral agudo o cortamente apiculado; 
lobulos 16-18(-21) x 9-13(-17) mm, obovados, relativamente extendidos, ligera 
e irregularmente reflexos marginalmente; estambres insertos en la base de la 
parte superior del tubo, anteras 6-7,4 mm de largo, glabras dorsalmente, la base 
auriculada, con las auriculas subtruncadas, cabeza estigmatica 2,7-3 mm de 
largo; ovario 1,3-1,7 mm de largo, diminuta y moderadamente puberulento; 
nectario anular, leve a moderadamente pentalobulado, ca. la mitad (o 
ligeramente menos) de la longitud total del ovario. Foliculos 15,5-18 cm x 1,5- 
3,5mm, glabros, levemente moniliformes; semillas 7-7,5 mm de largo, diminuta 
y densamente hirsutulas, coma 1,4-2,1 cm de largo, canela-amarillo. 

Distribucion, habitat y ecologia.—Restringida al E de Venezuela (estado de 
Bolivar) y Guyana (region Potaro-Siparuni), donde crece en bosques enanos y 
ventosos, en elevaciones de (450-)950-2000 m. Especimenes con flores han sido 
recolectados en Mayo y Octubre. Especimenes con frutos fueron recolectados 
en Mayo. 

Ensudescripcion de M. subcarnosa Benth. var angustata, Steyermark (1967) 
menciono que esta diferia de la variedad tipica (la que ocurre en el S y SE de 
Venezuela, Guyana y el N de Brasil en el Estado de Roraima) por sus hojas con 
las bases “angostadas y cuneado-agudas” (vs. redondeadas u obtusas a 
subcordadas), asi como laminas foliares oblanceoladas u oblanceolado-oblongas 
(vs. oblongas). En forma general estos caracteres son persistentes, pero existen 
suficientes caracteres adicionales que justifican el elevar su estatus a nivel 
especifico. De esta manera, M. a neta se puede diferenciar de M. subcarnosa 
por sus hojas con las t 


uadas (vs dispuestas 


L 


MORALES 3 359 


perpendicularmente), separadas por 4-10 mm entre si (vs. 2-5 mm), asi como 
inflorescencias sésiles o subsésiles, con el pedtnculo de 1-3 mm de largo (vs. 
pedunculadas y con el pedtinculo de 12-45 mm), foliculos de 16-18 cm de largo 
(vs. 7-13,5cm) y semillas diminuta y densamente hirsutulas, conel dumente 
erecto a suberecto (vs. densamente tomentulosas y cone 


Especimenes examinados. VENEZUELA. Bolivar: Piar, cima del Amaruay-tepui, 10 May 1986 (£1), 
Liesner & Holst 20762 (MO, VEN); Ptari-tepui, cercanias del campamenteo Misia Kathy, 28 Oct 1944 
(fl), Steyermark 59473 (MO). GUYANA: Montanias Pakaraima, Aymatoi, 17 Oct 1981 (f{D, Maas et al. 
5801 (K, MO, U), Maas et al. 5824 (U, WAG); region Potaro-Siparuni, rio Blackwater, 22 May 1991 (f1), 
McDowell et al. 4895 (US, WAG). 


Mandevilla boliviensis (Hook. f.) Woodson, Ann. Missouri Bot. Gard. 20:716. 1933. 
Dipladenia boliviensis Hook. f., Bot. Mag. 95:t. 5783. 1869. T1po: BOLIVIA: Datos perdidos (£1), 

Pearce 708 (HOLOTIPO: K, foto, MO). 
Mandevilla bella (Pittier) Woodson, Ann. Missouri Bot. Gard. 20:717. 1933, syn. nov. Dipladenia 
bella Pittier, J. Wash. Acad. Sci. 21:141. 1931. Tipo: VENEZUELA. DISTRITO FEDERAL: Hacienda 

rto La Cruz, region costera, 2300 m, 28 Ago-4 Sep 1918 (f), Pittier 8108 (HOLOTIPO: US). 

La unica diferencia mencionada por Woodson (1933) para separar Mandevilla 
bella de M. boliviensis (Hook. FE.) Woodson fue la longitud y forma del apice de 
los lobulos de la corola, asi como distribucion geografica disyunta. Sin embargo, 
no existen diferencias adicionales en otros caracteres morfoldgicos en los 
especimenes tipo que permitan reconocer ambos taxones. En todo caso, la 
longitud de los lobulos es muy variable en el material de M. boliviensis 
disponible hoy en dia, lo que en conjuncion con la alta variacion en la longitud 
y tamanio de la corola presente en otras especies del subgénero Mandevilla (eg., 
M. pohliana (Stadelm.) A. Gentry), impide reconocer ambos taxones. Otros 
sinonimos de M. boliviensis fueron anteriormente propuestos por Morales (1995, 


98). 


Mandevilla tristis J.F. Morales, sp. nov. (Fig. 2). Tipo: VENEZUELA. TACHIRA: Uribante, S 
ase de Cerro El Morro, 21 Jun 1990 (f1, fr), Dorr et al. 7115 (HOLOTIPO: NY; foto, INB) 


A Mandevilla bracteata (Kunth) Kuntze, M. hirsuta (Rich) K., Schum. M. sagittarii Woodson et M. 
steyermarkii Woodson, cui similis, foliis sessilis vel subsessilis, petiolis 1-2 mm longis, et bracteis 


2-2 mm latis differt 
Liana; ramitas algo aplanadas cuando jovenes, teretes a subteretes con la edad, 
ligeramente huecas, densamente tomentulosas cuando jovenes, esparcida a 
moderadamente puberulentas con la edad, mas raramente glabrescentes; 
coléteres interpeciolares inconspicuos, hasta 0,9 mm de largo. Hojas opuestas; 
peciolos 1-2 mm de largo; laminas foliares 2-7,3 x 1,5-3,2(-3,7) cm, ovadas a 
ovado-elipticas, el Apice cortamente cuspidado, la base cordada, los coléteres 
irregularmente distribuidos a lo largo del nervio central, membranaceas, 
diminuta y esparcidamente papilado-puberulentas en la superficie adaxial, la 
pubescencia mas densa sobre el nervio central, diminuta y densamente 


360 BRIT.ORG/SIDA 22(1) 


F 


4mm 


2mm 


4mm 


5mm 


2cm 


2mm 


2mm 


2cm 


Fic. 2. Mandevilla tristis (A-H de Werff & Gonzdlez 5313, |NB;\—J de Dorr et al. 7715, NY).A. Ramita con inflorescencia. B. 
Detalle de la sapere adaxial de la a mostrando los coléteres a lo largo del nervio central. C. Caliz, pedicelo y 
bractea. D. Vista de la adaxial de un sépalo, mostrando # coléter. E. Tubo de un botén parcialmente abierto, 
mostrando la posicién i: ie anteras y el tubo basalmente g F, Antera, vista dorsal. G. Cabeza estigmatica. H. 
Nectario y ovario. 1. Foliculos.J. Semilla 


MORALES 3 361 


tomentosas en la superficie abaxial, el indumento secando glauco, no revolutas, 
venas secundarias y terciarias conspicuamente impresas en la superficie abaxial, 
usualmente solo las venas secundarias apenas visibles en la superficie adaxial. 
Inflorescencia usualmente mas larga que las hojas subyacentes, axilar, densa a 
moderadamente puberulentas, con muchas flores, pedunculo 4-16(-20) mm 
de largo, pedicelos 1-1,5 mm de largo, raramente algunos pocos alcanzando los 
3 mm de largo en fructificaci6n, bracteas 9-15 x 1,2-2 mm, linear-ovadas, 
subfoliaceas; sépalos 8,5-11 x 0,5-1 mm, linear-ovados, largamente acuminados 
apicalmente, el apice no reflexo, escariosos, densa a moderadamente 
puberulentos externamente, el coleter solitario, irregularmente laciniado 
apicalmente; corola infundibuliforme, amarilla, algo giboso, densamente a 
moderadamente hispidulosa externamente, sobretodo en botones florales, el 
tubo giboso basalmente, la parte inferior 13-18 x 2-3,6 mm, la parte superior 
14-16 mm de largo, conica, 10-13 mm de diametro en la fauce, el apice del bot6n 
floral cortamente apiculado; lobulos 8-12 x 7-11 mm, obovados, relativamente 
extendidos; estambres insertos en la base de la parte superior del tubo, anteras 
3 2. 4,1 mm de largo, glabras dorsalmente, la base auriculada, con las auriculas 

d ,cabeza estigmatica 1,6-1,9 mm de largo; ovario 1,3-1,5 mm de largo, 
inconspicua y diminutamente papilado-puberulento a glabrescente; nectario 
anular, moderada a profundamente pentalobulado, ca. la mitad de la longitud 
total del ovario. Foliculos 10,5-13,5 cm x 1-3,6 mm, densa a esparcidarnente 
hispidulos, conspicuamente moniliformes; semillas 7,5-8 mm de largo, glabras 
a glabrescentes, coma 1,7-2,4 cm de largo, canela. 

Distribucion, habitat y ecologia.—Endémica al estado del Tachira en el E 
de Venezuela, donde crece en formaciones de sabanas y vegetacion arbustiva 
asociada a afloramientos rocosos o arenosos, en elevaciones de 1100-1300 m. 
Especimenes con flores y frutos han sido recolectados en Junio, pero material 
con flores se conoce también de Julio. 


Especimenes examinados. VENEZUELA. Tachira: Uribante, entre Siberia y Pregonero, 1] Jul 1983 ({D), 
Werff & Gonzalez 5313 (INB, MO). 


Mandevilla tristis se puede confundir con M. bracteata (Kunth) Kuntze, M. 
hirsuta (Rich) K. Schum., M. sagittarii Woodson y M. steyermarkii Woodson, 
un grupo de taxones que se caracterizan por sus inflorescencias con bracteas 
foliaceas o subfolidceas y corolas infundibuliformes. Sin embargo, M. tristis 
puede distinguirse facilmente por sus hojas subsésiles (pero aparentando ser 
sésiles), con los peciolos inferiores a 2 mm de largo (vs. 4-60 mm de largo), 
inflorescencias con las bracteas florales mucho mas angostas (1,2-2 mm vs. 3- 
9 mm), flores subsésiles, con el pedicelo hasta 1,5 mm de longitud (vs. 2-8 mm 
de largo) y corolas con el tubo mas pequeno y con un diametro mas angosto en 
la fauce. 


362 BRIT.ORG/SIDA 22(1) 


SUBFAMILIA APOCYNOIDEAE 
Tribu Apocyneae 
Odontadenia killipii Woodson, Ann. Missouri Bot. Gard. 18:546-547. 1931. Tipo: 
PERU. LORETO: Iquitos, 26 Sep 1929 (f1), Killip & Smith 29847 (HOLOTIPO: MO; IsoTIPO: US). 

Odontadenia killipii es un taxon relativamente facil de reconocer por el tamano 
relativamente pequeno de sus sépalos, corolas y foliculos. Aunque ha sido 
confundida con la comun O. verrucosa (Willd. ex Roem. & Schult.) K. Schum. 
ex Markgr, O. killipii puede reconocerse por sus sépalos relativamente mas 
pequenos (2-4 mm vs. 3-9 mm), corolas glabras externamente (vs. 
esparcidamente puberulentas a glabrescentes), con la parte superior del tubo 
de la corola anchamente conica (vs. angostamente conica) y foliculos mucho 
mas pequenos (7,5-10,5 cm vs. (13-)19-30 cm) (Morales 1999). Odontadenia 
killipii era conocida del S de Colombia a través de Ecuador hasta Pert, laGuyana 
Francesa y Brasil, pero se reporta por primera vez para Venezuela, es el estado 
de Amazonas. 


Especimenes examinados. VENEZUELA. Territorio Federal Amazonas: Rio Negro, Serrania de 
Tapirapeco, May 1992 (fl), Martin 1909 (MO, PORT); Atabapo, Oct 1989 (f1), Velasco 730 (MO). 


Tribu Mesechiteae 


Mandevilla Mage (Mull. Arg.) Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 
9:87, fig. 2L. 1924. Heterothrix ea vase Mull. Arg., Observ. Bot. 164. 1871. Eriadenia 
obovata Miers, Apocyn. S. Amer. 117-118, t. 14 B. 1878, nom. superfl. Tipo: PERU. LORETO: cerca 
de Tarapoto, fecha perdida (£D, Spruce ae (HOLOTIPO: W, foto F neg. 31792; IsOTIPOs: BM, CGE, 
F (fragmento), G, foto F neg. 26855, K [2 laminas], P) 


Este taxon fue originalmente reportado por Clausnitzer (1968) para Venezuela 
(pero sin testigo citado). Posteriormente, Morillo (1995) trato este taxon como 
“Mandevilla sp. A.’ en el tratamiento de la Flora de la Guayana Venezolana, 
aunque hizo referencia que esta especie habia sido conocida como M. 
vanheurchii. El estudio del tipo, asi como de diversos especimenes no deja duda 
de que este taxon esta presente en Venezuela, donde se conoce de los Estados de 
Amazonas y Bolivar. En forma general, esta especie se puede confundir con M. 
subcarnosa (Benth.) Woodson, ya que ambas comparten varios caracteres tales 
como tallos algo angulados cuando jovenes, hojas con el indumento ausente o 
glabrescentes y no evidente, inflorescencias con bracteas escariosas, con 
pedicelos cortos y corolas infundibuliformes, pero M. vanheurckii se distingue 
por facilidad por sus laminas foliares con las venas secundarias y terciarias 
usualmente no evidentes. 

Especimenes examinados. VENEZUELA. Amazonas: Casiquiare, alrededores de Yavita, rio Temi, 6-19 
Jul 1969 (£1, fr), Bunting et al. 3736 (U); Rio Negro, SE de San Carlos de rio Negro, 12 Nov 1987 (£1), 
Liesner & Carnevali 23012 (MO, WIS), cerro de la Neblina, rio Yatua, 14 Nov 1957 (fl, fr), Maguire et 
al. 42069 (F, NY, US); cerro de la Neblina, rio Yatua, 1 Ene 1958 (fl), Maguire et al. 42570 (K, NY, U, 
VEN), carretera San Carlos-Solano, 3 Feb 1977 (£1), Morillo 5153 (VEN): Atures, cano Piedra, SE de 


MORALES 3 363 


Puerto Ayacucho, Sep 1989 (f1, fr), Sanoja et al. 2965 (MO, NY, FOR Bolen {alpen tepui, Feb 1948 
(f1), Phel ps etal. 384 (NY); S Sororopan tepul, 14 Nov 1944 (£1) 


SUBFAMILIA RAUVOLFIOIDEAE 


Tribu Alstonieae 

El género Aspidosperma (Rauvolfioideae, Alstonieae) se encuentra ampliamente 
distribuido desde el S de México hasta Bolivia, Paraguay y el N de Argentina, asi 
comoen las Antillas, donde esta representado por pocas especies. En forma gen- 
eral, es uno de los grupos de las Apocynaceae neotropicales con una de las mas 
altas variaciones intraespecificas, en cuantoa la forma de las hojas e indumento 
de partes vegetativas e inflorescencias. Debido a esto, un sin numero de formas, 
variedades y taxones han sido propuestos para albergar diferentes estados dentro 
del rango de variacion de una misma especie, provocando una divergencia no- 
table en cuanto al numero de taxones aceptados en el género. De esta manera, 
Woodson (1951), reconocio un total de 52 especies, mientras que la revision de 
Marcondes-Ferreira (datos sin publ.) acept6 un total de 34 taxones. Clausnitzer 
(1968) report6 un total de 16 taxones para Venezuela, mientras que Morillo (1995), 
estimo que ca. 30 especies estaban presentes en el pais. Ahora bien, en el 
transcurso de identificaci6n de material sin identificar depositado en el Mis- 
souri Botanical Garden (MO), dos taxones anteriormente no conocidos en Ven- 
ezuela, pertenecientes al subgénero Aspidosperma (Marcondes-Ferreira & 
Kinoshita 1996) fueron encontrados, ambos restringidos a la cuenca baja 
amazonica. 


Aspidosperma rigidum Rusby, Mem. New York Bot. Sak 7:323. 1927. Tipo: BO- 
_LA PAZ: rio Bopi, 12 Sep 1921 (£1), Rusby 593 


Aspidosperma jaunechense A. Gentry, Phytologia 47:98-99. 1980, syn. nov. Tipo: ECUADOR. Los 
Rios: Vinces, bosque de Jauneche, 26 Mar 1980 (f1), Dodson & Gentry 9920 (HOLOTIPO: MO; 
ISOTIPO: SEL) 

Aspidosperma rigidum pertenece a la seccion Rigida (Marcondes-Ferreira & 
Kinoshita 1996) y ahora se encuentra distribuida en Costa Rica, Colombia, Ven- 
ezuela, Ecuador, Pert, Bolivia y el NO de Brasil. En general, esta especie se puede 
reconocer con facilidad por la siguiente combinacion de caracteres: fuste prin- 
cipal conspicua e irregularmente acanalado, tallos jovenes con las yemas 
apicales sin catafilos, hojas de 4-11 cm de largo, elipticas y membranaceas, 
inflorescencias laterales, con los lobulos de la corola angostamente elipticos y 
foliculos hasta 7 cm de largo, dolabriformes, lisos externamente y con las 
semillas secando negras. De forma vegetativa, se puede confundir con facilidad 
con las especies de la seccion Excelsa (Marcondes-Ferreira & Kinoshita 1996), 
pero A. rigidum se puede reconocer por sus laminas ovado-acuminadas y venas 
secundarias arquedas y unidas a la siguiente en forma terminal, formando una 
vena terminal. 

En la descripcion de Aspidosperma jaunachense, Gentry (1980) tnicamente 


364 BRIT.ORG/SIDA 22(1) 


clasific6é ese taxon dentro de la clasificacion intragenérica propuesta por 
Woodson (1951), pero sin discutir en forma directa sus relaciones con alguna 
especie en particular. Sin embargo, aunque él cité que este taxon era muy simi- 
lara A. rigidum, no profundiz6 en discutir las diferencias que podrian separar 
ambos taxones. De hecho, el estudio de los tipos ha revelado que no existen 
diferencias significativas que justifiquen la validez de A. jaunachense por lo que 
es reducido a la sinonimia. 


Especimenes examinados. VENEZUELA. Territorio Federal Amazonas: mision rio Mavaca, || Feb 1991 
(El, fr), Stergios et al. 15335 (INB, MO, PORT) 


Aspidosperma schultesii Woodson, Ann. Missouri Bot. Gard. 38: 168, tl, f.6. 1951. 


} 


Tipo: BRA SIL: A | Ira i-Igarapeé ) V bit atluente 


del rio Taraira, 4-6 Jul 1948 ({1), Schultes & Lopez 10178 (HOLOTIPO: MO: ISOTIPO: F). 


Aspidosperma schultesii, pertenece a la seccion Schultesia (Marcondes-Ferreira 
& Kinoshita 1996), y dentro del género es un taxon bastante distintivo que se 
puede reconocer con facilidad por sus ramitas jovenes sin catafilos, tallos y 
peciolos secando negros, glabrescentes, laminas foliares con la venacion 
secundaria apenas evidente 0 no evidente, asi como corolas con los lobulos ca. 3 
veces mas largos que el tubo, y foliculos negros al secar, con la superficie externa 
conspicuamente granulada. Esta especie se encuentra ahora distribuida en Co- 
lombia, Venezuela, N de Brasil y Peru. 


Es pees examinados. VENEZUELA. Territorio Federal Amazonas: San Carlos de Rio Negr 


al de 
la conf] lel Rio Negro y el brazo Casiquiare, 29 Feb 1980 (fr), Clark & Maquirino 7367 (MO). 


AGRADECIMIENTOS 


Los siguientes herbarios facilitaron el préstamo o uso de sus colecciones: BM, 
CGE, FE K, MO, NY, P, PORT, U, US, W, WIS, VEN. Quiero reconocer la ayuda 
prestada por Gerardo Aymard (PORT) por su disposiciOn a suministrar una 
referencia bibliografica especifica. Asimismo, agradezco al Missouri Botanical 
Garden (MO) por el apoyo brindado que ha hecho posible la visita de su herbario 
y por la cantidad de especimenes enviados como regalo por identificacién y a 
Alina Freire-Fierro y Rosa Ortiz-Gentry, por su colaboracién en mis visitas a 
dicho herbario. Finalmente, se agradece la revision y comentarios por parte de 
David Goyder y un revisor anonimo 


REFERENCIAS 


A.vorce, L. 1985. Monographie des Apocynacées—Tabernaemontanoidées Américaines. 
Mém. Mus. Natl. Hist. Nat., B, Bot.30:1-216 
CLAUSNITZER, |. 1968. Los géneros de Apocynaceae de Venezuela. Acta Biol. Venez. 3: 
4 


Enoress, M. & PV. Bruyns. 2000 A revised classification of the Apocynaceae s.|. Bot. Rev. 66: 


MORALES 3 365 


LeeuwenserG, A.J.M.1994.A revision of Tabernaemontana two.The New World species and 
Stemmadenia. Royal Botanic Gardens, Kew, U.K. Pp. 213-450. 

Marconbes-F erreira, W.& L. KINOSHITA. 1996.Uma nova divisdo infragenérica para Aspidosperma 
Mart. (Apocynaceae). Revista Brasil. Bot. 19:203-214. 

Markarar. F. 1971 [1972]. New Apocynaceae and Asclepiadaceae from Venezuela. Acta 
Biol. Venez. 6(1-4):65—74 

Markarar. F. 1978. Novedades de Apocynaceae. Acta Biol. Venez. 13:353-355. 

MonacHino, J.1958.Apocynaceae. In:B. Maguire y J. Wurdack, eds. The botany of the Guyana 
Highland Ill. Mem. New York Bot. Gard. 10:117-138. 

MonacHino, J. 1961.Apocynaceae.In:B. Maguire y J.Wurdack, eds. The botany of the Guyana 
Highland lV.Mem. New York Bot. Gard. 10:58-65. 

Morates, J.F. 1995. An evaluation of the Mandevilla boliviensis complex. Phytologia 78: 
197-198. 

Mora es, J.F.1998.A synopsis of the genus Mandevilla (Apocynaceae) in Mexico and Cen- 
tral America. Brittonia 50:214-233. 

Morates, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae). In: AJ.M. 
Leeuwenberg, ed. Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67: 
381-477. 

Monrates, J.F.2005.Estudios en las Apocynaceae Neotropicales X:definicién de la verdadera 
identidad de Mandevilla leptophylla, con la descripcién de dos nuevas especies y una 
nueva combinacion de Mandevilla (Apocynoideae, Mesechiteae) para Sur América. 
Sida 21:1537-1548. 

Moritto, G. 1989a. Las Apocynaceae del estado Lara. Biollania 6:193-237. 

Moritto,G.1989 b. Notas preliminares sobre el género Mandevilla Lindley (Asclepiadaceae) 
en Venezuela. Ernstia 53:15—20. 

Monritto, G. 1995. Aspidose In:J.A.Steyermark, P Berry y B. Holst, eds. Fl. Venez. Guayana 
2:481-490. Timber Press, Portland, OR and Missouri Botanical Garden, St. Louis. 

Moritto, G. 1995 Mandevilla. In: J.A. Steyermark, P. Berry y B. Holst, eds. Fl. Venez. Guayana 
2:518-529. Timber Press, Portland, OR and Missouri Botanical Garden, St. Louis. 

STEYERMARK, J. 1967. Flora del Auyan-Tepui. Acta Biol. Venez. 2(5-8):5-370. 

Woopson, R.E. 1933. Studies en the Apocynaceae IV. The American genera of Echitoideae 
XXVI. Ann. Missouri Bot. Gard. 20:605-790. 

Woooson, R.E. 1948. Miscellaneous new Apocynaceae and Asclepiadaceae. Ann. Missouri 
Bot. Gard. 35:233-237. 

Woopson, R.E. 1951. An interim revision of the genus Aspidosperma Mart. & Zucc. Ann. 
Missouri Bot. Gard. 38:1 19-206. 

ZARUCCHI, J.,G.N. Moritto, M.E. Enpress, B.F. HANSON & A.J.M. LEeUWwENBERG. 1995. Apocynaceae. In: 
JA. Steyermark, P Berry & B. Holst, eds. Fl. Venez. Guayana 2:471—571.Timber Press, Port- 
land, OR and Missouri Botanical Garden, St. Louis. 


366 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Davip STUART. 2004. Dangerous Gardens: The quest for plants to change our lives. 
(ISBN 0-674-01104-X, hbk.). Harvard Press, Cambridge, MA (Orders: TriLiteral 
LLc, 100 Maple Ridge Dr, Cumberland, RI 02864-1769, U.S.A., 800-405-1619 
http://www.hup.harvard.edu/). $35.00, 208 pp., color drawings, color and 
b&w photo, bibliography, and index, 7 1/2" x 10 3/8" 


Do poisonous plants and aphrodisiacs interest you? What about medicinal plants, life elixirs, and 
“mind expanding” plants? Then you should venture into the enjoyable new book called Dangerous 
Gardens: The i st for plants to change our lives. 

David Stuart has created this intriguing text about mankind's use of plants to pursue 


h, haps and long life. The book is divided into chapters that focus on different subject ar- 

eas to which plants have been see by humankind: “Great Afflictions,” “ The Vital Organs,” “The 
Flight a Pain,” “Chasing Venus,” “The Killing Plants,” “The Seven Ages of Man,” “The Mind” and 
“Mysteries of the Gods.” The author ee both historical background and historical reasoning 
regarding why plants were ana for different ance or desires. For example, the book begins witha 
history of the pl Other oan ve through- 


~_ 


healt 


out the book ne now plants were aed oe used for one reason and accidentally or eventu- 
ally learned that the same plant actually cured other illnesses. Foxglove (Digitalis dela for 
example, in the 1600/1700s was used for wounds and lung ailments; not until the late 1700s was it 
learned that this same plant had dramatic effects on the heart. The text also describes many ae 
that were liberally used in history until their true and often-dangerous behavior was unearthed. 
One example is ee w mach up until he late ee was used liber oF as a cure for au kinds of 


ailments. The I th { famous 
poisoners, “cure-all” quack pills, ee use in rituals ad the use of various - plants as aphrodisiacs. 
The author also included information in s chapter what oS pas labelled "shadow plants,” 

plants that are often used for treatment, sucl ] f| pp.), but that science 


and/or the medical community a not ceed the true benefit and risks, if any. 

I found Dangerous Gardens hard to put down, it reads like a very good cable TV beumenay 
on plant use by man, complete with photographs and/or drawings of the plants being discussed, 
practitioners of herbal medicine, perpetrators of poisoning, quackery, and many other historical 


images related to plant use. This book was insightful regarding the rationale that was used to decide 
which Cea to use for different illnesses. It was fascinating to learn about how various plants were 
used diff y throughout time. This book is recommended to anyone with an interest in the topic 


of plant use by mankind; it is easy to read and engaging.—Lee Luckeydoo, Herbarium, Botanical Re- 
search Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(1): 366. 2006 


TAXONOMIC OVERVIEW OF THE HETEROTHECA 
VILLOSA COMPLEX (ASTERACEAE: ASTEREAE) 


Guy L.Nesom 


Botanical Research Institute of Texas 
t 
Fort Worth, Texas 76102-4060, U.S.A. 


ABSTRACT 


ieterthecs villosa (as treated by senile 1996, 2006) is a comipiex species with nine varieties, most 
of Heterotheca villosa var. nana and H. villosa var. scabra are 
ena silonarrc and intergrade Tittle, but each is widely sympatric with H. villosa and distinct 
from it. Recognition at specific rank accurately reflects the status of var. nana and var. scabra, and 
they are treated here, respectively, as Heterotheca horrida (Rydb.) Harms and Heterotheca polothrix 
Nesahy nom. et stat. nov. pieleoiiees senoptiila sensu stricto is distinct from H. stenophylla var. 


g with it,and the latter is appropriate mere villosa 
Pele oe Fe (Rx741 


var. angustifolia ydb.) Harms. The New Mexico endemic Heterotheca vil 


osa var. sierrablancensis 
Semple is raised to specific rank as Heterotheca sierrablancensis Sane Ny ,comb et stat. nov. 
Identifications of vars. villosa, foliosa, ballardii,and minor (all judge- 


ents because of their broad sympatry and extensive intergradation. The disimeten between var, 
el nculata and H. zionensis is not clear, and both taxa apparently intergrade broadly with more 
typical H. villosa. Variety depressa is maintained at specific rank as H. depressa (Rydb.) Dorn. Maps 
show the generalized distributions of the taxa of the H. villosa complex sensu Semple, and a nomen- 
clatural summary outlines an alternative taxonomy. 


RESUMEN 
Heterotheca villosa (as treated by sae e 1996, 2006) pl ho variedades 
la mayoria de las cuales son sim varios eee Hehe rothec villosa var. nana y ie L villosa 
P g 
] ae 
A oO ye i 
simpatrica con H. villosa y dik le ella. El imi ivel ifi flej 


yi 
el estatus de var. nana y var. scabra, y son tratadas aqui Gonieleiete ol ecelion r ida 


(Rydb.) Harms y Heterotheca po 


othrix tstat. nov. Heterot! 

es diferente de H. stenophylla var. ene ean Semple) y simpatrica con ai, y esta ultima es 

euiada Apropcamien( como H. Valo var. sangusyol ia aay Harms. ea pcicemies de Neevo 
villosa var. 

sierrablancensis (Semple) Neco comb. et stat. nov. Las identificaciones ae las vars. villosa, foliosa, 


pl picen at bido a su amplia simpatria e 


ballardii, y minor (todas sensu 


intergradacion extensa. La diferenci tre var. pedunculata y H. zionensis no es clara, y ambos 

axa aparentemente se intergradan con la mas tipica H. villosa. La variedad depressa se mantiene en 
el rango especifico como H. depressa (Rydb.) Dorn. En los mapas se muestran las distribuciones gen- 
erales de los taxa del cas H. villosa sensu Semple, y un resumen nomenclatural delinea una 
taxonomia alternativa 


Intermediates between taxa of Heterotl ften 1 and unequivo- 
cal identifications are correspondingly difficult. Phenotypic plasticity, small 


SIDA 22(1): 367 — 380. 2006 


368 BRIT.ORG/SIDA 22(1) 


degrees of differentiation, and hybridization apparently underlie the taxonomic 
difficulties. Tetraploids are common and gene exchange among them appar- 
ently occurs frequently. A monograph of Heterotheca sect. Phyllotheca (Semple 
1996) provides a detailed view of the taxonomic structure of this group and 
addresses difficulties in identification 

Heterotheca villosa, as treated by Semple (1996, 2006), is a widespread and 
complex species with nine varietal taxa, most of which are geographically over- 
lapping or almost completely superimposed. Typical H. villosa (var. villosa) oc- 
cupies essentially the northern half of the species range, but it is broadly sym- 
patric with var. minor, var. foliosa, var. ballardii, and var. nana. In part of 
Nebraska and South Dakota, five varieties (sensu Semple) of H. villosa occur 
sympatrically; four varieties occur sympatrically in Saskatchewan; three oc- 
cur sympatrically in much of Wyoming, Colorado, and Utah. Two are relatively 
narrow endemics (within a single state); all the others occur in at least four 
states. All taxa were mapped in detail by Semple (1996, Figs. 39 and 40); gener- 
alized outline maps shown here (Figs. 1, 2, 3) are derived in most part from 
Semple’s dot maps. 

Regarding the Heterotheca villosa complex, Semple (1996, p. 114) noted that 
it might be “logical to merge all ... varieties together with no infraspecilic taxa 
being recognized ... This would result in the loss from the formal nomenclature 
of a great deal of information on variation and distribution in what is admit- 
tedly a difficult species complex.” His approach (as noted by Nesom 1997) has 
been to recognize sympatric conspecific varieties, using formal varietal names 
to identify what many (or most) systematists might regard as geographical 
trends or populational variants. “A variety is characterized by all members of a 
population exhibiting a particular morphology distinct from other individu- 
als in the species. The distribution of these populations is sympatric with popu- 
lations whose members are not within the same variety, and also many popu- 
lations of morphological intermediates exist [citing various references]. .. Van 
Steenis described varietal level variation as being continuous with other vari- 
eties, although the continuum would have pronounced modes” (Semple 1974, 
p. 8-9). Similarly, and with regard to Heterotheca villosa, “the races fit well with 
the concept of variety in that each occurs in pure form in some populations, 
and the overall ranges are sympatric to a considerable degree with at least one 
other variety” (Semple 1996, p. 108). 

Semple has used a similar taxonomic approach for Heterotheca sessiliflora 
(Nutt.) Shinners sensu lato, a primarily Californian complex, and formal 
changes in taxonomic rank were proposed by Nesom (1997) to clarify its sys- 
tematics. The taxonomic situation with H. villosa (Pursh) Shinners is consid- 
ered here. Harms (1968, 1970) also has addressed aspects of the taxonomy of 
the H. villosa complex. 


NESOM, TAAUNUMIC OVERVIEW MI 369 


| ae 


® 

+ 
Fic. 1. Generalized distribution of Heterotheca villosa (A) var. villosa and (B) var. minor, both Semple, and (C) H 
villosa var.an gustifolia. Illustrated is the broad sympatry of I {th | sympatry/alloy 
try of var. angustifolia. The range of H. villosa d | minor, includ I N da (Clark Co., Excelsior 
Canyon, 7 Sep 1941, Clokey 8759-MO).Th ge of ifolia includ I lowa (Lyon Co., prairie, among 


Sioux Quartzite exposures, nw corner of county, 25 Aug 1924, Shimek s.n.-MOQ). 


370 BRIT.ORG/SIDA 22(1) 


Fic. 2. Generalized distribution of Heterotheca villosa (A) var. foliosa, (B) var. ballardii, (C) H. zionensis, and (D) var. 
pedunculata, all sensu Semple. Compare with Fig. 1 to see broad sympatry of var. foliosa and var. ballardii with var. 


villosa and var. minor. 


Status of Heterotheca villosa var. pedunculata 

Heterotheca villosa var. pedunculata occurs in complete sympatry with H. villosa 
var. minor in the Four Corners region, as indicated by Semple (1996) (var. 
pedunculata with upper cauline leaves densely to extremely densely strigose, 
thus pale green to white, eglandular to very sparsely glandular; 2n = 18, 36; var. 
minor with upper cauline leaves sparsely to moderately densely strigose, 
sparsely to densely glandular; 2n = 18, 36). Variety minor “is the most variable 
linfraspecific taxon] in the species and includes some local distinctive 
morphotypes that grade into other forms. Plants intermediate between this and 
all other varieties occur in areas where the ranges are sympatric, and they make 
infraspecific taxonomy of the species difficult” (Semple 2006). In a study of 


NESOM, TAXONOMIC VVYECRVICVY VI 371 


~ \ A ian 
bet 


Fic. 3.G lized distributi f (A) Heterott | lothrix, (B) H. horrida, (C) H. stenophylla (D) H. depressa and (E) H. 
£4 2 Pes + bE L 


sierrablancensis. Ill ha dictrihut; 


3 


Four Corners Heterotheca, | have been able to make only arbitrary morphologi- 
cal distinctions between var. pedunculata and var. minor. 

Variety pedunculata is very similar to Heterotheca zionensis Semple and 
partly sympatric with it. Plants of H. zionensis are said to be identified by stems 
and leaves appearing silvery to whitish from densely strigose vestiture and pe- 
duncles and phyllaries usually densely glandular but otherwise glabrate. Such 
plants are distinguished by Semple from more densely pubescent individuals 
of H. villosa var. minor or from plants identified as var. pedunculata (with gray 
to silvery leaves) by glandular peduncles and phyllaries, even though plants of 
var. minor may have glandular phyllaries and populations of H. zionensis may 


372 BRIT.ORG/SIDA 22(1) 


include individuals with eglandular peduncles and phyllaries—“non-glandu- 
lar forms of H. zionensis occur in north-central Utah; glandular and non-glan- 
dular forms occur in the Utah-Arizona border region” (Semple 1987, p. 385). The 
geographic range of H. zionensis overlays the range of the western portion of 
var. minor as well as of var. pedunculata, and distinctions among these are un- 
clear. It does seem clear that silvery-leaved plants are concentrated in the south- 
west part of the range of H. villosa, but if the silvery-leaved plants are represen- 
tative of a geographic trend in vestiture, widely intergrading with more typical 
plants, then it is likely that the names H. zionensis and H. villosa var. 
pedunculata represent plants shaped by the same trend and the same underly- 


ing genetics. 


Status of Heterotheca villosa var. nana and var. scabra 

Heterotheca villosa var. nana (diploid) is almost completely overlaid in its geo- 
graphic range by var. foliosa (diploid and tetraploid but tetraploid in its area of 
overlap with var. nana, many chromosome counts, fide Semple 1996) and by 
var. minor (diploid and tetraploid, many counts, fide Semple 1996). Variety nana 
varies considerably and probably hybridizes with H. villosa and H. fulcrata 
(Greene) Shinners (Semple 2006), but even so, var. nana is morphologically con- 
sistent and recognizable throughout its range and has previously been treated 
at specific rank by various botanists as H. horrida (Rydb.) Harms (e.g., Harms 
1968, 1970; Great Plains Flora Association 1986; Dorn 1988). 

Heterotheca villosa var. scabra is the westernmost taxon treated by Semple 
within H. villosa, except for the south-reaching extension of the species in Cali- 
fornia (var. minor fide Semple). Semple (1992) noted that var. scabra is closely 
related to the Californian endemic H. shevochii (Semple) Semple (originally 
described as H. villosa var. shevochii). It seems equally plausible that var. nana 
is closely related to var. scabra. All three taxa tend to have spreading leaves and 
strongly developed glandularity. 

The geographic ranges of var. nana and var. scabra are slightly overlap- 
ping along the Utah/Colorado and northern Arizona/New Mexico borders (Fig. 
3). Semple (2006) observed that “In the Four Corners area, var. nana grades into 
var. scabra, and collections often are difficult to place into one of the two taxa”— 
but I find the distinction clearer and intermediates not numerous, especially in 
view of the broad range and relative morphological constancy of var. scabra. 
Plants of var. scabra and “aff. var. scabra” cited by Semple at a southeastern ex- 
tension of its range in trans-Pecos Texas are tentatively identified here as forms 
of H. viscida (A. Gray) Harms. New Mexico collections of var. scabra well east of 
the range illustrated in Fig. 3 (San Miguel Co., Rio Arriba Co. cited but not 
mapped by Semple 1992) are better identified as var. nana. | have not seen records 
of var. scabra from Idaho or Colorado (as indicated on Fig. 3), but it is a distine- 
tive entity and Dr. Semple’s citations document its presence there. 


— 


NESOM, TAXONOMIC VYLAVICW UE 373 


Variety nana and var. scabra are distinguished by Semple by various dif- 
ferences (mostly foliar), and my study corroborates and emphasizes their dis- 
tinction, as in the following couplet. 


1. Cauline leaves usually not contorted upon drying, upper mostly lanceolate to tri- 
angular-lanceolate, acute at apex; leaf surfaces moderately to densely glandular, 
the glands ue elie ~~ the sparsely strigose (nonglandular) vestiture, 
nonglandular hai t ricted to midvein area (middle 1/3) of adaxial surfaces 
or sometimes nearly absent on both surfaces, especially on more distal leaves; phyl- 
laries glabrous to very sparsely strigose, glandular; 2n = 18, 36 (many counts, fide 
Semple 1996) Heterotheca villosa var. scabra 
Cauline leaves often contorted (margins undulate) upon drying, upper mostly ob- 
long or oblong-lanceolate, acute to obtuse at apex; leaf surfaces moderately to 
gosely Senauiay me glands conspievous a the ee hispid to 
lyt telyd ie 
Goce. glandular; ae = 18 (many counts; 2n = 27, one count, ide Semple 1996) 
eter Shee villosa var. nana 


Including some features more difficult to precisely contrast, plants of var. scabra 
(in contrast to var. nana) have stems and leaves with much reduced non-glan- 
dular vestiture but with denser and more conspicuous glandularity, leaves dif- 
ferently shaped, smaller, and more widely spaced, heads relatively fewer, and 
phyllaries glabrous (lacking non-glandular hairs) but glandular. The two taxa 
are different in geography. 

In their area of sympatry, var. scabra is distinguished from a regional, more 
typical form of Heterotheca villosa (var. minor fide Semple) by the following 
contrasts. 


1. Upper cauline leaves lanceolate to triangular-lanceolate, acute at apex; leaf sur- 

faces moderately to densely glandular, the glands little obscured by the sparsely 

Sstrigose (nonglandular) vestiture, nonglandular hairs often restricted to midvein 

area (middle 1/3) of adaxial surfaces or sometimes nearly absent on both surfaces; 

phyllaries glabrous to very sparsely strigose Heterotheca villosa var. scabra 
Upper cauline leaves linear to oblanceolate, oblong-lanceolate, or narro 

vate, acute to obtuse at apex; leaf surfaces eglandular or sparsely to cones glan- 

dular, moderately to densely strigose evenly over both surfaces; phyllaries sparsely 

strigose Heterotheca villosa var. minor 


— 


In view of their morphological integrity and apparent reproductive isolation 
from elements of more nearly typical Heterotheca villosa, both H. villosa var. 
nana and H. villosa var. scabra are treated here at specific rank. 


Heterotheca polothrix Nesom, nom. et stat. nov. Chrysopsis villosa var. scabra Eastwood, 
roc. Calif. Acad. Sci. 2.6:294. 1896. Heterotheca villosa var. scabra (Eastwood) Semple 
Phytologia 73:453. 1992. Type: U.S.A. UTAH. San Juan Co. Willow Creek, 14 Jul 1895, A eee 
38 (HOLOTYPE: CAS, digital image!). Non Heterotheca scabra DC. 1836. The species is named 
for a resemblance of the medial strip of adaxial leaf vestiture to the mane of a pony (Greek, 
polos, pony, and thrix, hair). Blake's varietal epithet (‘cinerascens,’ at varietal rank, as below) 
is poorly descriptive of the taxon (as also noted by Semple 1987). 


374 BRIT.ORG/SIDA 22(1) 


Chrysopsis viscida A. Gray subsp. cinerascens Blake, Proc. Biol. Soc. Washington 35:173. 1922. 
Heterotheca horrida subsp. cinerascens (Blake) Semple, Brittonia 39:381. 1987. TYPE: U.S.A. 
UTAH. [Beaver Co. fide Welsh 1982] Beaver Creek, among rocks in the oak region, 2 Sep 1901, I. 
Tidestrom 2873 (HOLOTYPE: US, internet image). 


Heterotheca horrida (Rydb.) Harms, Wrightia 4:17. 1968. Chrysopsis horrida Rydb, 
Bull. Torrey Bot. Club 31:648. 1905. Type: U.S.A. COLORADO. |Weld Co. New Windsor, 8 Aug 
1900, G.E. Osterhout 2326 (HOLOTYPE: NY internet image! ISOTYPES: RM, WIS). 
Chrysopsis canescens var. nana A. Gray, Mem. Amer. Acad. Sci. 4[PI. Fend1.:78. 1849. Heterotheca 
villosa var. nana (A. Gray) Semple, Novon 4:54. 1994. TyPE: U.S.A. NEW MEXICO. [Mora Co.?!: 
elevated rocky region 2 mi E of the Mora River, Aug 1847, A. Fendler 391¢ (HOLOTYPE: GH). 


Status of Heterotheca villosa var. sierrablancensis 

Heterotheca villosa var. sierrablancensis was described by Semple (1996) from 
the White Mountains and Sacramento Mountains of Lincoln and Otero cos., 
New Mexico. It is similar to H. horrida in its conspicuously glandular vestiture 
and congested and consistently corymbiform capitulescence but distinct in its 
leaves more densely strigose, the upper cauline ascending, larger, and differ- 
ently shaped (lanceolate or oblong-lanceolate to ovate), ray corollas longer, and 
habitats at higher elevation. It occurs at the southeastern corner of the range of 
H. horrida (H. villosa var. nana) and Semple indicated (1996, Fig. 3) that the two 
taxa are most closely related to each other. Both are primarily diploids. The ex- 
tended population system of var. sierrablancensis is morphologically variable 
but it lies inside the geographic range of H. horrida and apparently is isolated 
from it as well as the regional expression of H. villosa, both of which occur at 
lower elevations. Recognition at specific rank is appropriate for this taxon, es- 
pecially in view of the parallel rank of H. horrida. 


Heterotheca sierrablancensis (Semple) Nesom, comb. & stat. nov. Heterotheca vil 
var. sierrablancensis Semple, Univ. Waterloo Biol. Ser. 37:146. 1996. TYPE: U.S.A. NEW MEXICO. 
Lincoln Sierra Blanca, NM 532 8.5 km W of NM 48, rocky outcrop and face of roadcut at 

shback directly below scenic overlook, 2 Oct 1995, J.C. Semple and Semple 10513-A (Ho- 
LOTYPE: WAT, ISOTYPES: MO!, NMC, RM). 


OSd 


Status of Heterotheca villosa var. depressa. 

Semple treated this narrowly endemic taxon at varietal rank because it hybrid- 
izes with var. minor. Both taxa are tetraploid in their area of sympatry. As noted 
by Semple (2006), “Var. depressa grows in geyser basins in Yellowstone National 
Park and is locally common in rocky soils in Teton National Park; it is very rare 
farther south in Wyoming. It is distinguished by its small stature, small heads, 
and linear-oblanceolate leaves with many hairs and usually some glands. It is 
similar to some forms of var. minor, which occur throughout the range of var. 
depressa and with which it hybridizes. ... Typical var. depressa occupies the vi- 
cinity of hot springs and geyser basins and nearby river flood banks, while var. 
minor occupies other drier or non-geyserite nearby habitats.” Markow (2001, 


NESOM, TAXONOMIC VVYCAVICYY UT 375 


2004) found that the occurrence of H. depressa is strongly correlated with that 
of another narrow endemic, Stephanomeria fluminea Gottlieb. 

The geographic range of Heterotheca villosa var. depressa is relatively dis- 
crete, like that of H. sierrablancensis, and is essentially imbedded within the 
much larger distribution of H. villosa sensu lato. Although var. minor (sensu 
Semple) hybridizes with var. depressa, the latter appears to have its unique eco- 
logical niche, maintains its morphological identity, and is justifiably treated at 


specific rank. 

Heterotheca depressa (Rydb.) Dorn, Vasc. Pl. Wyoming (ed. L) 295. 1988. Chrysopsis 
depressa Rydb., Mem. New York Bot. Gard. 1:381. 1900. Heterotheca villosa var. depressa (Rydb.) 
Semple, Novon 4:53. 1994. TyPE: U.S.A. WYOMING. Teton Co.: Yellowstone National Park, Lower 
Geyser Basin, 7000 ft, 4 Aug 1897, PA. Rydberg and E.A. Bessey 5067 (HOLOTYPE: NY internet 
imagel; ISOTYPES: CAN, NY, US internet image!). 


— 


Status of Heterotheca stenophylla var. angustifolia. 

Semple (1996, 2006) has treated Heterotheca stenophylla (A. Gray) Shinners as 
comprising two varieties, var. stenophylla and var. angustifolia (Rydb.) Semple, 
“based on field studies and the results of multivariate analyses .... The type of 
var. angustifolia is morphologically closer to many indi Is of [H. stenophylla] 
var. stenophylla than it is to either H. canescens or typical H. villosa. ... At the 
very least, the type of var. angustifolia belongs in H. stenophylla, regardless of 
where the more canescens-like and the more villosa-like plants are placed” 
(Semple 1996, p. 94). 

Previous botanists have treated var. angustifolia within Heterotheca villosa 
(e.g., Harms 1968, 1970; Great Plains Flora Association 1986). Harms observed 
(1968, p. 16-17) that “over most its range, including Texas, Heterotheca 
stenophylla appears to represent a quite distinct diploid (n = 9) species, but some 
tetraploid (n = 18) populations are known from a narrow band along the east- 
ern edge of the species’ overall distribution reaching from south-central Kan- 
sas tothe Wichita M tains of south-central Oklahoma (to ?Archer Co., Texas) 
which hybridize and intergrade somewhat with [tetraploid] H. villosa. Asa re- 
sult, the tetraploid H. stenophylla populations as a whole are usually distinguish- 
able from the diploid populations by the presence of such introgressed charac- 
ters from H. villosa as more pubescent involucres, stems, or leaves, but it seem 
impossible to assign individual plants to a particular ploidy level on this basis. 
Thus it appears impractical to attempt to taxonomically distinguish the tetra- 
ploid from the diploid H. stenophylla populations despite the the presence of 
an obvious reproductive barrier based upon different ploidy levels. Neither does 
it seem logical to taxonomically merge H. stenophylla with H. villosa [var. 
angustifolia] merely because certain tetraploid populations of the former hy- 
bridize more or less extensively with the latter, when most of the H. stenophylla 
taxon is diploid and clearly distinct morphologically and biologically.” 


376 BRIT.ORG/SIDA 22(1) 


Heterotheca stenophylla (sensu stricto) is broadly sympatric with var. 
angustifolia and I (with Harms) also find that the two are distinct in the field 
and herbarium. Triploid putative hybrids have been reported (Semple 1996, 
2006). In contrast, var. angustifolia intergrades with broader-leaved forms of H. 
villosa, especially in Nebraska and the Dakotas, and is reasonably treated within 
H. villosa. “Inclusion of var. angustifolia ... only slightly increases the morpho- 
logical complexity of H. villosa, as defined by Semple, and [it] occupies a part of 
the overall geographic range es its sympatry with conspecific varieties is 
relatively less” (Nesom 1997, 

Harms (1970, p. 1567) digea “a ‘most of the Texas material [of Heterotheca 
villosa] belongs to var. foliosa (characterized by dense canescent appressed pu- 
bescence of involucres and herbage, and larger heads) or to var. angustifolia 
(characterized by narrower leaves, coarser and sparser herbage pubescence and 
smaller heads); these varieties intergrade on a broad scale, and much Texas 
material appears to be more or less intermediate between them.” Still, a trend 
within H. villosa toward narrow leaves is pronounced in Texas, Oklahoma, and 
Kansas, and identification of these plants as var. angustifolia is justifiable. 


i 


. Stems eglandular; leaves oblong-oblanceolate to elliptic-oblanceolate, 3-6 mm 

wide, eglandular or sessile-glandular on abaxial surface, moderately to densely stri- 

gose, hairs usually without inflated basal celss; phyllaries eglandular or inconspicu- 

ously and minutely glandular; chromosome counts of 2n = 36 (fide Semple 1996) 
Heterotheca villosa var. angustifolia 
. Stems sessile- oe leaves linear to narrowly ol oblanceolate, 2-3(—4.5) mm wide, 

sessile-glandular, sparsely hispid-strigose, hairs with conspicuously inflated and vis 

cid basal =alle aS eile glandular to eglandular, sparsely strigose or essentially 

without nonglandular hairs; many chromosome counts of 2n = 18, few of 2n= 

(fide Semple 1996) Heterotheca stenophylla 


aaa 


Consistent distinction between Heterotheca villosa var. angustifolia and H. 
canescens (DC.) Shinners is sometimes more difficult than between var. 
angustifolia and H. stenophylla. Heterotheca canescens is mostly diploid, but 
some tetraploids have been reported, and as noted by Semple (1996, p. 97), 
“tetraploids [of H. canescens and H. villosa var. angustifolia] apparently hybrid- 
ize forming local swarms of parent-like and hybrid individuals.” And “in Texas, 
it sometimes is difficult to clearly distinguish herbarium specimens of [H. 
canescens] from certain canescent, strigose-sericeous, smaller-headed, more 
narrow-leaved forms of H. villosa (Harms 1970, p. 


Heterotheca villosa (Pursh) Shinners var. angustifolia (Rydb.) Harms, Wrightia 

16. 1968. Chrysopsis a Rydb., Bull. Torrey Bot. Club 37:128. 1910. Chrysopsis 

villosa var. angustifolia (Rydb.) Crongq., Bull. Torrey Bot. Club 74:150. 1947. Heterotheca 

senophla vat. iis See (Ryd) Semple, Novon 4:53. 1994. Type: (Semple 1990): U.S.A. 

NEBRASKA. Hooker Co.: Middle Loup River, near Mullen, on sandhills, 14 Sep 1893, PA. Rydberg 
1766 (LECTOTYPE: NY internet image! ISOLECTOTYPES: GH, NY, US). 


NESOM, TAXONOMIC OVERVIEW OF 377 


ee stenophylla (A. Gray) Shinners, ao & Lab. 19:68. 1951. Chrysopsis 

var. stenophylla A. Gray, Boston J. Nat. Hist. 6 (PL. Lindh.):223. 1850. Chrysopsis villosa 

var. epee (A. Gray) A. Gray, Synopt. FI. : Amer. (ed. 2) 1(2):123. 1884. ees 

stenophylla (A. Gray) Greene, Erythea 2:96. 1894. Type: U.S.A. TEXAS. [Llano Co, Llano, from 

strong ligneous roots in crevices of smooth granite rocks, Nov 1847, EJ. Lindheimer 631 
LOTYPE: GH; ISOTYPES: CAN, DS, GH, K, MO!, ND-G, NYI4 sheets, internet image], US). 


Taxonomic summary of the Heterotheca villosa complex 

Four taxa treated by Semple as varieties within Hetorotheca villosa are recog- 
nized here at specific rank: H. horrida, H. polothrix, H. sierrablancensis, and H. 
depressa. Intergradation and sympatry among var. villosa, var. foliosa, var. 
ballardii,and var. minor make it impractical and biologically unrealistic to rec- 
ognize them as more thana single entity. The identity of var. pedunculata vs. H. 
zionensis and vs. other densely hairy plants in the region of var. minor is not 
clear. Variety angustifolia appears to be justifiably recognized within H. villosa 
(vs. within H. stenophylla) 

Distinct morpl hic entities are recognized here with formal names. 
Relatively stable morphology and sympatry with close relatives indicate that 
such taxa are reproductively isolated, and they are treated at specific rank. In 
addition to typical Heterotheca villosa, var. pedunculata (sensu lato) and var. 
angustifolia appear to be geographically distinct but each intergrades with more 
typical plants where ranges meet—these latter two are tentatively regarded here 
at varietal rank. It might be useful to recognize widespread morphological 
“nodes” within var. villosa sensu lato at the rank of forma or with an informal 
name; the contrasting taxonomic approach (formal recognition of sympatric 
conspecific varieties) requires a majority of typological and arbitrary judgments. 

Local and regional floristic studies may encounter and identity stable sym- 
patric races that call for a more complex nomenclature than proposed here. But 
because of the geographic mix of hybrids and ploidy levels, biological and 
morphological patterns in the Heterotheca villosa complex may remain more 
complex than can be adequately circumscribed by traditional nomenclature 


_ 


eK 


1. Heterotheca villosa (Pursh) Shinners [1951 
Amellus villosus Pursh (1814, NEOTYPE: Missouri River, almost certainly North Dakota] 


la. Heterotheca villosa var. villosa 


Including laa, lab,and lac. The sympatry and intergradation of var. villosa sensu 
Semple with var. foliosa, var. ballardii, and var. minor suggest that these taxa 
are arbitrarily identified. 

[laa]. Heterotheca villosa var. iosa (Nutt.) Harms [1968] 

Chrysopsis foliosa Nutt. (1841, TYPE: sw WYOMING] 

Chrysopsis' villosa var. see iosd ut ) ssa [1947] 

Chrysop: ata Is. [190 PE: COLORADO. El Paso/ Teller Co.] 

eee ae var. ae 7 oe Nels. [1909 


378 BRIT.ORG/SIDA 22(1) 


If Chrysopsis foliosa var. imbricata is treated as synonymous with Heterotheca 
villosa var. foliosa (as done by Semple 1996, 2006), the earliest name at varietal 
rank is “var. imbricata.” In this context, H. villosa var. foliosa is an invalid name. 


[lab]. Heterotheca villosa var. ballardii (Rydb.) Semple [1994] 

Chrysopsis ballardii Rydb.[1931, TYPE: MINNESOTA. Carver Co] 
“This is the generally robust, larger-headed, many-rayed, eglandular, oblong- 
leaved race of the species occurring on the northeastern prairies of Canada and 
the United States” (Semple 1994, p. 53). “Forms of var. ballardii are sufficiently 
similar to forms of var. foliosa that the two taxa might be treated as convarietal 
races under the name var. foliosa” (Semple 1996, p. 114). The geographic range 
of var. ballardii lies within that of var. villosa sensu Semple. 


[lac]. Heterotheca villosa var. minor (Hook.) Semple [1994] 

Chrysopsis villosa var. minor t look. [1847, TYPE: WYOMING. Sweetwater Co] 
Diplopappus hispidus Hook eee TYPE: SASKATCHEWAN. Carlton House] 
Ch rysopsis villosa var. hispida (Hook.) A. Gray [1884] 

Heterotheca villosa var. hispida (Hook.) Harms [1974] 


Canadian botanists have commonly recognized two varieties of H. villosa, var. 
villosa (leaves and involucres with appressed, nonglandular vestiture) and var. 
hispida (leaves and involucres with spreading hairs intermixed with glands), 
but none has indicated that the taxa are geographically or ecologically distinct 
or that intergradation and typological identifications are not prevalent in this 


~~ 


region. 
1b. Heterotheca villosa var. pedunculata (Greene) Harms ex Semple [1987] 


Chrysopsis pedunculata G [1900, TYPE: COLORADO: Archeluta Co.] 
?Heterotheca zionensis Sem ple [1987. TYPE: UTAH. Washington Co]] 


Representing the densely strigose, silvery-sericeous expression of the H. villosa 
complex; common in Utah, northern Arizona, and the Four Corners region. As 
noted above, I have not been able to distinguish H. zionensis from var. 
pedunculata, based on criteria established by Semple. 
1c. Heterotheca villosa var. angustifolia (Rydb.) Harms [1968] 

Chrysopsis angustifolia Rydb. cea TYPE: NEBRASKA. Hooker Co] 

Chrysopsis villosa var. angustifolia (Rydb.) Crongq. [194 

Heterotheca stenophylla var. quantal (Rydb.) Semple [1994] 


Allopatric and distinct from other expressions of H. villosa, at least in Texas, 
Oklahoma, and most of Kansas, intergrading in Nebraska, South Dakota, and 
North Dakota. 
2. Heterotheca horrida (Rydb.) Harms [1968] 

Chrysopsis horrida Rydb. [1904, TyPE: COLORADO. Weld Co.] 


Heterotheca villosa var. nana (A. Gray) Semple [1994] 
Chrysopsis canescens var. nana A. Gray [1849, TyPE: NEW MEXICO, ?Mora Co. 


NESOM, TAXONOMIC OVERVIEW OF MPLEX 379 


Ww 


. Heterotheca sierrablancensis (Semple) Nesom [2006] 


Heterotheca villosa var. sierrablancensis Semple [1996, TYPE: NEW MEXICO. Lincoln Co. 


> 


. Heterotheca polothrix Nesom [2006] 


Chrysopsis villosa var. scabra Eastwood [1896, TyPE: UTAH. San Juan Co] 
Heterotheca villosa var. scabra (Eastwood) Semple [1992 
. Heterotheca depressa (Rydb.) Dorn [1988] 
Chrysopsis depressa Rydb.[1900, TYPE: WYOMING. Teton Co|] 
Heterotheca villosa var. depressa (Rydb.) Semple [1994] 


WN 


ACKNOWLEDGMENTS 


All pertinent specimens at BRIT/SMU, MO, and TEX/LL were studied, and a 
loan from SJNM was helpful in providing a base of study for plants of the Four 
Corners region. In all, more than 1350 specimens of the Heterotheca villosa com- 
plex were examined. Additionally, I have examined all collections of all taxa of 
Heterotheca at these herbaria. Staff at CAS provided an image of the holotype 
of Chrysopsis viscida subsp. cinerascens and made observations of details of its 
vestiture. Robert George lent his digital proficiency toward production of the 
maps. Constructive review comments of Vernon Harms and an anonymous 
reviewer are greatly appreciated. 


Postcript.—Dr. John Semple has read some or all of the manuscript, and in any 
case, he is intensely opposed to the approach and its conclusions. But it has not 
seemed necessary to re-assemble and re-cite the massive number of specimens 
(10,300) Dr. Semple had on hand for his monographic study of sect. Phyllotheca. 
He has noted that his treatment is based on multivariate analyses (yet unpub- 
lished), and no statistics are found here (nor do I believe that taxonomic con- 
clusions based on such an approach must necessarily be correct). I do not have 
the long and focused field experience he has had with this group. Still, because 
of our strong and basic differences in taxonomic philosophy (regarding con- 
cepts of species and varieties) and apparently because of differences in the way 
in which we perceive and interpret patterns of variation, it obviously is pos- 
sible to differ in taxonomic conclusions regarding the same group of plants. We 
agree on the delimitation of some of the taxa but not others. For those we agree 
should be formally recognized, we disagree at what rank some should be treated. 
If this taxonomic overview is seen as no more than a “superficial opinion-piece,” 
at least I have tried to provide a clear rationale for the basis of my opinions and 
why they differ from those of Dr. Semple. The alternative taxonomy proposed 
here allows me to identify plants of the Heterotheca villosa complex in a con- 
sistent and biologically meaningful way, from my point of view. Others also 
have a choice. 


380 BRIT.ORG/SIDA 22(1) 


REFERENCES 


Dorn, R.D. 1988.Vascular plants of Wyoming (ed. 1). Mountain West Publishing, Cheyenne, 
Wyoming. 

GREAT PLAINS FLorA Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. 

Harms, V.L. 1968. Nomenclatural changes and taxonomic notes on Heterotheca, including 
Chrysopsis,in Texas and adjacent states. Wrightia 4:8-20. 

Harms, V.L. 1970. Heterotheca. In: D.S. Correll and M.C. Johnston. Manual of the vascular 
plants of Texas. Texas Research Foundation, Renner, Texas. Pp. 1563-1569. 

Markow, S. 2001.Report on a survey for Stepbhanomeria fluminea in Grand Teton National 
Park. Prepared for the Greater Yellowstone Network Inventory and Grand Teton Na- 
tional Park. Wyoming Natural Diversity Database. <http://uwadmnweb.uwyo.edu/ 
wyndd/ Reports/pdf_markow/Stephanomeria_Markow01.pdf> Accessed February 
2006. 

Markow, S. 2004. Survey for Stephanomeria fluminea on the Bridger-Teton National Forest. 
Prepared for the Bridger-Teton National Forest ae the BealOe 4 is Service Office. 
Wyoming Natural Diversity Database. <http:/ ndd/R 
pdf_markow/ Stephanomeria_Markow04.pdf> Accessed Fabris 2006. 

Nesom, G.L. 1997. Review: “A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms 
(Compositae: Astereae)" by J.C. Semple. Phytologia 83:7-21. 

SemPte, J.C. 1974. The phytogeography and systematics of Xanthisma texanum DC.: proper 
usage of infraspecific categories. Rhodora 76:1-19. 


Sempte, J.C. 1987. New names, combinations, and lectotypifications in Heterotheca 
(Compositae: Astereae). Brittonia 39:379-386 


Sempte, J.C. 1990. Neotypification of Amellus villosus, the identity of a Bradbury collection, 
and typification of some other goldenasters (Compositae: Astereae). Brittonia 42: 
22) 5228. 

Sempte, J.C. 1992. The goldenasters of California, Heterotheca (Compositae: Astereae 
names and combinations. Phytologia 73:449-455, 

SemPte, J.C. 1994. New combinations in the Heterotheca villosa (Pursh) Shinners complex 
(Compositae: Astereae). Novon 4:53-54. 

Sempte, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: 
Astereae). Univ. Waterloo Biol. Ser. 37:i-iv, 1-164 

Sempe, J.C. 2006. Heterotheca (Asteraceae: Astereae). In: Flora of North America Editorial 
Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford Uni- 
versity Press, New York and Oxford. Vol. 20. 

WetsH, S.L. 1982. Utah plant types——historical perspective 1840 to 1981-—annotated list, 
and bibliography. Great Basin Naturalist 42:129-195, 


WS 


new 


TAXONOMY OF LANTANA SECT. LANTANA 
(VERBENACEAE): 1 CORRECT APPLICATION OF 
LANIANA CAMARA AND ASSOCIATED NAMES 


Roger W. Sanders 
Botanical Research Institute of Texas 
09 Pecan Street 
Fort Worth, Texas 76102-4060, U.S.A. 
rsanders@brit.org 
ABSTRACT 


The previous lectotypification of Lantana camara L. is evaluated by examination of characters of 
the lectotype, review of oo original material, and documentation of current usage of the name. 

The current usage is analyzed by surveying Peranieat acu and by eeu specimens for an- 
notations between 1753 and the q t f those specimens. Current 
usage of L. camara includes a eadely cultivated and naturalized cultigen species of hybrid origin 


that is taxonomically distinct from L. camara. To determine the correct name of the cultigen, al 
names in Lantana’ sect. Lant d, typil Hes if TISCESSAE YE and Bessils and son amucaMy 


disposed. h il i I | is 
K W. Sanders. ace of the oaea De brids in the L.camara ee is hypothesized. oa new com- 
p. aculeata (L S R. me paneets and Lantana nivea Vent. Subse. 


mutabilis (WJ. Hook ) R.W. Sanders. Nineteen | 


RESUMEN 


La lectotipificacion previa de Lantana camara L. se evaltia mediante examen de los caracteres del 
lectotipo, revision de otro material original, y documentacion del uso actual del nombre. El uso ac- 
tual se analizé revisando la bibliografia adecuada y por muestreo de especimenes entre 1753 y el 

nte por comentarios en relacion a caracteres criticos de esos especimenes. El uso actual de L. 
y naturalizada de origen hibrido que es 


pres 
camara incluye una especie ampliamente cultivada 
smi dif L. camara. Para determit el nominee correcto ne la planta cultivada 


los | bre de Lantanasect. Lantana osible, 
y taxonomicamente decidido. No hay nombres disponibies as . mana eu a nas se describe 


como Lantana strigocamara R.W. Sanders. Se hace una 


en el complejo L. camara. Se hacen dos an ees ok Daven camara subsp. aculeata (L.) 
R.W. Sanders y Lantana nivea Vent. subsp. mutabilis (WJ. Hook.) R.W. Sanders. Se realizan diez y nueve 
lectotipificaciones y siete epitipificaciones. 

Plants that have gone under the Linnaean name Lantana camara L. are well 
known, not only as hardy summer ornamentals worldwide but also as natural- 
ized weeds having devastating economic impact in humid tropical areas of the 
Old World (Wolfson & Solomons 1964; Howard 1970; Stirton 1977; Swarbrick 
et al. 1995; Day et al. 2003). The systematics of these cultivated and naturalized 
plants is not well understood, even though considerable effort has been made 
to elucidate their biological control (Day et al. 2003). This confusion arises in 


SIDA 22(1): 381 — 421. 2006 


382 BRIT.ORG/SIDA 22(1) 


part from the convoluted history of exploration, cultivation, hybridization, and 
artificial selection that began at least 60 years before the publication of Linnaeus’ 
Species Plantarum (1753). Records that have been compiled (Howard 1969; Stirton 
1977) suggest that, during the eighteenth century, fanciers hybridized different 
wild species and infraspecific taxa of Lantana L. sect. Lantana from Mexico, the 
West Indies, and Brazil. As shown in this paper, the early validly published names 
were based on garden-grown material (almost entirely so until 1817 and com- 
monly so into the 1850s), much of which consisted of hybrid combinations. 

As herbarium specimens of wild-collected species of sect. Lantana became 
available after the early 1800s, many of the available, poorly distinguished 
names were frequently misapplied to them. Adding to this confusion was the 
introduction of cultivated hybrids into neotropical regions where indigenous 
taxa occur. Due to the propensity of lantanas to undergo polyploidy and the 
partial fertility of odd polyploid levels (Natarajan & Ahuja 1957; Khoshoo & 
Mahal 1967; Spies 1983, 1984; Spies & Stirton 1982a, b, c; Sanders 1987a, b), even 
more complex hybrids formed between indigenous taxa and the escaped hy- 
brid cultigens (Sanders 1987a, b, c, 1989a). Thus, the limits of natural variation 
have been obscured, a the ability of taxonomists to develop effective 
classifications for the grou 

Schauer (1847), Br ae (1895), and Troncoso (1974) developed current sec- 
tional concepts. Species of Lantana sect. Lantana (=sect. Camara Cham.) are 
characterized by predominantly narrow floral bracts that are somewhat in- 
conspicuous among the tubular corolla bases, by usually yellow or orange pig- 
mented corollas (white-flowered populations known in several species [unpubl. 
observ. see also discussion below concerning purplish pigments in the section), 
and by blackish drupes. Each drupe contains a characteristically inflated com- 
pound endocarp that resembles a horse’s skull in which the seed chambers are 
in the position of the eye sockets. Some of the species of the other major section, 
Lantana sect. CallioreasCham., might be confused with those of sect. Lantana. 
Generally, species of sect. Callioreas are described as involucrate, usually hav- 
ing ovate to reniform conspicuously imbricate floral bracts and purplish co- 
rollas. However, fruits are needed to insure correct sectional placement—drupes 
are usually white or purplish and endocarps are subglobose, bilobed (seed 
chambers fill each hemisphere), noninflated, and reticulately ornamented. 

Schauer (1847) published the only worldwide revision of Lantana in De 
Candolle’s Prodromus. All more recent work has been limited to regional and 
garden floras. The foremost student of Verbenaceae in the twentieth century, 
Harold N. Moldenke, never produced a revision of Lantana, but he did describe 
a number of new species and infraspecific taxa. Horticulturalists, ecologists, 
and some floristic taxonomists have effectively submerged much of Lantana 
sect. Lantana into L. camara, treating this wide spectrum of variation asa single 


SANDERS, TAXONOMY OF LANTANA 383 


species, (e.g., Kuntze 1891; Troncoso 1965, 1974; Bailey Hortorium 1976; Schemske 
1976, Huxley et al. 1992; Cullen et al. 2000). 

Sanders (1987a, b, c, 1989a, b) undertook a study of the variation of wild 
and naturalized populations of Lantana sect. Lantana in Florida and parts of 
the West Indies. By correlating chromosome numbers and meiotic behavior with 
morphology, he was able to distinguish the natural (often diploid) wild taxa 
from the naturalized and spontaneous hybrids and to develop morphological 
taxonomic criteria. Sanders found that plant architecture, leaf and bract size 
and shape, indument features, inflorescence/infructescence development and 
size, fruiting bract persistence, and floral pigment classes and dosage effects 
correlate well with cytology, geographic distributions, and sf delimitations. 
On the other hand, he argued that development of prickles and specific corolla 
colors, characters relied upon by many other authors, vary too widely within 
and among taxa to be effective criteria, a conclusion recently supported by 
molecular studies (Scott et al. 1997; Day et al. 2003). Further support comes from 
the work of Isidro Méndez S. of Cuba who took up the study of Cuban and West 
Indian Verbenaceae (1992, 1993, 2002). Méndez explicitly accepted the taxo- 
nomic criteria proposed by Sanders for Lantana sect. Lantana. 

The natural taxa of sect. Lantana fall into three separate phenetic groups 
(to be validated in a subsequent paper; cladistic status not yet determined) on 
the basis of trichome structure on the abaxial surface of the leaf blades, as well 
as shape, adaxial luster, and venation of the leaf blades. The “pilose-morph” 
species (Pilose Group, Fig. 1) are cl terized by soft, filiform, spreading (erect 
or curly) hairs usually densely disposed on all veins and intervening tissue. Most 
of these have leaf blades that are broadly ovate to narrowly deltate, rounded to 
cordate at the base and acuminate to obtuse at the apex, dull above (epidermis 
poorly reflective), and more or less pinninerved. The “strigose-morph” species 
(Strigose Group, Fig. 2) have the abaxial hairs developed as conspicuous strigae, 
i.e. the hairs are stout, conical, antrorsely geniculate, and are scattered and re- 
stricted to the midrib, and secondary, tertiary, and sometimes higher order veins, 
but not to the intervening tissue. Leaf blades are usually ovate-elliptic, often 
tapering at base and apex, lustrous above, and more or less triplinerved. The 
“setose-morph” species (Setose Group, Fig. 3) abaxially bear well-separated, long, 
flexible to subrigid, spreading, setiform trichomes that are more or less restricted 
to the midrib and secondary and tertiary veins. Otherwise they are like the pi- 
lose-morph species. Thus, even though all three groups have adaxial strigae, 
the Stigose Group is the only one with strigae on both surfaces. 

Despite Méndez’s (2002) acceptance of Sanders’ criteria, he disagreed with 
Sanders over the application of the name L. camara. Because L. camara is the 
proposed type of the genus (Jarvis 1992; Jarvis et al. 1993) and one of the two 
oldest names in the secton, the correct application of most other names in the 


BRIT.ORG/SIDA 22 


= 


a 
LM 


ey 
oy | 
=! 


Fic. t nen, paises INE ie A), ee Group. a. inflorescence. b. rey tative leaf blad 


section hinge upon the correct application of L. camara. Asserting that Sanders 
had committed two errors, Méndez stated: 


Sanders (1989a]) attributed this binomial [L. camara] to a phenotype very different from prevailing 
opinion among the authors later to Linnaeus and not in agreement with the lectotype selected by 
Moldenke & Moldenke (1983) 

However, until the present study, analysis of the lectotype and subsequent ap- 
plication of the name have not been straightforward. The critical characters 
were not visible on the lectotype since all of its leaves were mounted adaxial 
side up. Sanders (unpubl.) was not able to determine whether the lectotype 
matched the pilose-morph or the strigose-morph plants, especially given the 
quality of the microfiche and xerographic images available at the time. How- 
ever, because H. N. Moldenke had selected the type, Sanders (unpubl.) relied 
primarily on the numerous annotations of Dr. Moldenke (Table 1, App. B, see 
also Moldenke 1980a) to develop a concept of the species and apply the name. 
Sanders’ concept was further reinforced by annotations of C.D. Adams, and the 


1) 


SANDERS, TAXONOMY OF LANTANA 385 


Fic. 2.Lantana scabrida,| (Soland ?,,BM) ing Stri G a.infl { 


ae £. L on | £ £ 


c Scale bars:a = 1cm;b&c=1mm. 


leaves. b. 


treatments of Lantana in The Flowering Plants of Jamaica (Adams 1972) and A 
Flora of Tropical Florida (Long & Lakela 1971), as well as on identified culti- 
vated material received under that name on the U.S. National Arboretum, 
Longwood Gardens, and tabl | nurseries. In all cases, the plants 
were characterized by leaves dominated by strigose-morph hairs, cordate-ovate 
blades, and dull upper surfaces! (here subsequently called the “Strigose-Cor- 
date-Dull-” or “SCD cultigen”, Fig. 4). In the West Indies, some wild-collected 
specimens bear codominant mixtures of both hair morphs. However, cytologi- 
cal study confirmed these to be hybrids between the SCD cultigen and native 
taxa of the Pilose Group (Sanders 1987b). Thus, Sanders concluded that the lec- 
totype had the strigose-morph hairs, and he applied the name L. camara spe- 
cifically tothe SCD cultigen. Many of the described species in the Pilose Group 
(including L. horrida, L. tiliifolia,and L. arida) he lumped into a single species 
and applied the name L. urticifolia Mill. 

Méndez appears to have studied more recent, higher quality images in con- 
nection with a project to lectotypify Linnaean species of Verbenaceae of the 
West Indies in collaboration with Steve Cafferty of the Linnaean Typification 
Project (Méndez & Cafferty 2001). Although he, too, was unable to see the abaxial 
surfaces, he concluded on other grounds that the lectotype matched the pilose- 


'Adams included in his concept of L.camara the SCD cultigen and L. scabrida. The few specimens of L. scabrida 


that | studied and bear his annotation as L. camara actually have lustrous upper leaf surfaces. 


386 BRIT.ORG/SIDA 22(1) 


al 
al 


Fic. 3.1 hii (Hi) 20499, TEX) ing $ a.inflorescence.b 


J t 


leaf surface. Scale bars:a=5 mm;b&c=1mm. 


morph, not the strigose-morph plants. He applied the name L. camara to the 
taxon to which Sanders applied the name L. urticifolia. 

If Méndez’s two assertions merit redress, then another, later-published name 
should be applied to the SCD cultigen that Moldenke, Adams, and Sanders called 
Lantana camara. Of course, current usage, regardless of usage by authors of the 
early post-Linnaean period, must be established. If current usage of L. camara 
has clearly replaced the Linnaean concept with the concept of the SCD culti- 


gen, then conservation of the nomenclatural type could serve as an alternative 


solution. 

Therefore the purpose here is to |) evaluate the lectotypification by review- 
ing its publication, by clarifying the characters of the lectotype, by understand- 
ing current usage of L. camara, and by determining any variance between the 
type and current usage; 2) affirm the correct application of that name, 3) if nec- 
essary, determine which other name is to be applied to the SCD cultigen, and 4) 
dispose of all other names, known to me, that are applicable to the complex. 


Typification of Lantana cama 

Moldenke and Moldenke (1983) effectively lectotypified Lantana camara by 
citing LINN 783.4 as “type” (ICBN, Art. 7.11, Greuter et al. 2000). This 
lectotypification was accepted by Sanders 1989b) and Méndez (2002). In March, 
2004, | visited BM, and Dr. Charles Jarvis, long-time participant in the Linnaean 
Typification Project, consented to break the lower stem to permit study of the 
abaxial surface of the lowest two leaves (Fig. 1). The type, indeed, is of the pi- 
lose-morph and is verified by leaf shape and inflorescence structure as an ele- 
ment of the naturally occurring species distributed from the Bahamas and 
Greater Antilles, through Mexico south to northwestern South America. Thus 
of Méndez’s two assertions, the one that “Sanders (1989) attributed this bino- 
mial toa phenotype...not in agreement with the lectotype selected by Moldenke 
and Moldenke (1983)” is correct. 


Taste 1.Data documenting history of annotation by H.N.Moldenke and other taxonomists familiar with the classification of Lantana."v.moritz” indicates plants 
annotated as L. camara var. moritziana. See text for further discussion. 


Years Authorities N Pilose % Mixed % Strigose % Setose % 
1981-2005 Moldenke (incl. v. moritz) 62 15 24% 20 32% 21 44% 0) 0% 
Moldenke (excl. v. moritz) 47 0 0% 20 43% 27 57% 0 0% 
tner 45 ¢) 0% 25 56% 20 44% @) 0% 
All (incl. v. moritz) 107 15 14% 45 42% 47 44% 0 0% 
All (excl. v. moritz) 92 0 0% 45 A9% 47 51% 0) 0% 
1956-1980 Moldenke (incl. v. moritz) 153 ifs) 8% 56 37% 83 54% i 1% 
Moldenke (excl. v. moritz) 14] ] 1% 56 40% 83 59% ] 1% 
Other 60 0 0% 4] 68% 19 32% 0 0% 
All (incl. v. moritz) 213 13 6% 97 46% 102 48% ] 0% 
All (excl. v. moritz) 201 ] 0% 97 48% 102 51% ] 0% 
1931-1955 Moldenke 16 2 13% 6 38% 7 44% ] 6% 
Other ] 0 0% 0 0% ] 100% @ 0% 
All 17 2 12% 6 35% 8 47% ] 6% 
1906-1930 All 16 0 0% 10 62% 6 38% 0 0% 
1881-1905 All 4 0) 0% 3 75% ] 25% Q 0% 
1856-1880 All 2 0 0% 0 0% 2 100% 0 0% 
1831-1855 All 3 1 33% ] 33% ] 33% 0 0% 
1753-1830 All 15 10 67% 2 13% 1 7% 2 13% 


VNVINV1°L35 VNVLNVT JO AWONOXVL‘SUIONYS 


L8€ 


388 BRIT.ORG/SIDA 22(1) 


b. inflorescence 


Development of current usage 

Linnaeus’ concept of L.camara was developed from an array of cultivated and 
horticulturally selected plants. The synonyms and illustrations that he cited 
in the protologue are based on vouchers that are primarily elements of the Pi- 
lose Group. All Hortus Cliffortianus (1737) specimens are either the same spe- 
cies as LINN 783.4 (319 Lantana 1-B!, Linn. Herb. specimen at S [a cut-down 
Herb. Cliff. specimen, C. Jarvis, pers. comm., dig. photo!]), are hybrids between 
that taxon and the Strigose Group (319 Lantana I), or are elements of L. horrida 


SANDERS LANTANA SECT. LANTANA 389 


Kunth (319 Lantana 1-C), 1-D!, and 320 Lantana lol[the latter possibly hybrid- 
ized with the Strigose Group]). Moreover, an unnumbered Herb. Cliff. sheet (!) is 
an element of the Strigose Group (L. splendens Medik.). 

Linnaeus did not see the vouchers for the syntype illustrations. One of these 
(icon in Plukenet, Phytographia 385. t. 114, f. 4. 1691. [Voucher: 98:143 top-left 
specimen, BM-SL!]), belongs to the Setose Group (L. hirsuta M. Martens & 
Galeotti). The Commelin syntype has no known voucher (C. Jarvis, pers.comm.), 
illustrates only an inflorescence, and cannot be placed to trichome-morph. Even 
so, the trichome characters may not have been of much concern to Linnaeus. 
For example, in 1767, he did segregate L. mista, which has distinctly hispid twigs, 
as depicted in a Dillenius plate (see App. B), which he cited. However, Linnaeus 
did not use this character to differentiate the new species, but rather used the 
protracted leaf-blades, longer bracts, and capitula with mixed corolla colors. 
According to C. Jarvis (pers. comm.), no other original material is known. 

To determine how this somewhat broad Linnaean concept was modified 
by later botanists, I present two lines of evidence. The first isa survey of sampled 
specimens to determine annotation patterns by taxonomists (Table 1, Fig. 5). 
The specimens are all those annotated by H. N. Moldenke as L. camara in LL/ 
TEX (including the Moldenke Herb.) and BRIT/SMU, a selection of those at K, 
and a few for which | obtained photographs at BM, LINN, and OXE Besides 
Moldenke, annotating authorities include: Linnaeus, Medikus (implied by cita- 
tion, 1775), Schauer, Urban, Merrill, Hutchinson, R. Meikle, J. K. Morton, R. 
Fernandes, B. Verdcourt, G. Bromley, and S. Atkins. Dr Méndez and I are excluded. 

Plants annotated as L. camara (or in a few cases as L. aculeata, see App. B) 
were scored for whether trichomes on the abaxial surface were pilose-morph 
only, strigose-morph only, setose-morph only, or codominant mixtures of stri- 
gose- and pilose-morphs (including rare mixtures of strigose- and setose-mor- 
phs). Between the Linnaean period, when L. camara was applied primarily to 
pilose-morph plants, and the late nineteenth century, specialists began to ap- 
ply the name primarily to the strigose-morph plants or those with mixed tri- 
chome morphs. For those annotated specimens from the neotropics, the mixed 
trichome plants are presumed hybrids primarily between the naturalized SCD 
cultigen and native species of the Pilose Group. In the paleotropics, the mixed- 
trichome plants appeared to be either 1) hybrids between escaped pilose-morph 
plants and the SCD cultigen, 2) escaped cultivars selected from hybrids between 
L. camara and various species of the Strigose Group, or 3) naturalized plants 
consisting of complex spontaneous hybrids between plants of the preceding 
two categories. 

As this annotation trend developed, the plants of the Pilose Group began 
to be annotated with other later names. In particular, note the quotation of H. 
Moldenke [apparently in personal correspondence] by Howard (1969): 


390 BRIT.ORG/SIDA 22(1) 


BO% 
70% 

‘ | 
wn 60% a ! 
| a} . ra x | 
3 \ ig . | 
hen * 
~ 50% s < | 
seed ? ee -_=- * * = 
2 : oe. Ren ee =A 
£ nad 4 fa 
<z 40% "aiid Salen Aaa a u 
rs] \? x 
e 30% + 
E , 

O 20% + 
e 
\ t) 
See a i 


PNAS 
. * Saeed 
0% — alana >o- 


T 


1763-1830 1831-1905 1906-1930 1931-1955 1956-1980 1981-2005 


Year 
—e—Pilose - @ - Mixed —a — Strigose — - Setose 
—e—Pilose +mrz - & - Mixed+mrz —tz, — Strigose + mrz 


Fic. 5. Plot of portion of data from Table 1, showing historical pattern of specialists’ application of the name Lantana 
camara to pilose-morph taxa (mostly L. camara, some L. horrida and their hybrids) vs. strigose-morph taxa (mostly L. 
strigocamara, L. nivea, and L. sanslouel vs, oe eibiibab the two groups une morph plants) vs. setose-morph 


plants (mostly L. hirsuta). “+ mrz f L. camara var. moritziana. 
You accuracy of identification on the labels of plant a marked ‘Lan- 
tana camara’ in ne | have found that a large percentage of such material is actually L. 


moritziana, L.glandulosissima, L. scorta, L. horrida, L. arida, L. glutinosa, etc. etc 


In other words, authorities (e.g... Moldenke and others) attributed specimens of 
L. camara to other species names referable to the Pilose Group. Surprisingly, in 
the mid 1970s, Dr. Moldenke accepted Lopez-Palacios’ (1974) reduction in rank 
of L. moritziana asa variety of L. camara. If that anomaly is excluded from the 
analysis, then the application of L. camara to strictly pilose-morph plants by 
taxonomic specialists is negligible over the last 50 years. 

The second line of evidence is a survey of the literature (App. A). Adequate 
descriptions of leaf indument are scarce prior to the 1850s, but those by Medikus 
(1775), Sprengel (1825), and Schauer (1847, 1851) all indicate a prevailing accep- 
tance of plants with the abaxial surfaces soft-hairy. The first publication di- 
verging and treating strigose-morph plants as L. camara is by Otto and Dietrich 
(1841). This reflected a growing confusion as to what L. camara really is. By the 


SANDERS, TAXONOMY OF LANTANA 391 


beginning of the twentieth century, most horticultural taxonomists took a broad 
concept of L. camara to include both pilose- and strigose-morph (or even se- 
tose-morph) plants in the species. This sentiment is reflected in Bailey’s Cyclo- 
pedia (1900) 

Lantanas have been long in cultivation, and it is difficult to refer the garden f botanical spe- 
cies. The species themselves are confusing. Most of the garden kinds are of the L. Car mara type. There 


are several Camara-like species which probably have pbudized to produce these forms; but Voss 
] 


(1896], the latest garden monographer, regards these s] forms of L. Camara (preferring, 


however, to use the name L. aculeata). Accepting L. Camara in Voss’s sense, the garden lantanas may 


be said to be derived from that species; and this view is adopted below. 


Over the last 100 years, most floristic taxonomists (and hence, ecologists) have 
developed a similar attitude and broad concept or have followed Moldenke 
(1970) and Adams (1972), applying L. camara to the SCD cultigen and its hy- 
brids, as has Sanders. Therefore, Mendez’s other assertion that “Sanders (1989) 
attributed this binomial toa phenotype very different from prevailing opinion 
among the authors later to Linnaeus” is inaccurate. 


Disparity between the lectotype and current use 

The disparity between the lectotype and current use can be viewed in two ways. 
Either the current usage 1) includes the type and encompasses both pilose- and 
strigose-morph plants, or 2) excludes the type and is applied only to the SCD 
cultigen (and its hybrids with other species). The first view is problematic be- 
cause the SCD cultigen behaves as a species distinct from the species named L. 
camara by Linnaeus. The second view raises the issue of relectotypifying L. 
camara to fit current usage. This might be preferable if LINN 783.4 were a hor- 
ticultural hybrid no longer extant. However, the type is an element of a wide- 
spread wild species to which the name L. camara has been properly applied, at 
least in part, by taxonomists Standley (1924), Leon & Alain (1957), Macbride 
(1960), Gibson (1970), Nash & Nee (1984), Lopez-Palacios (1977), Méndez (2002), 
and many ecologists. Futhermore, neither original material nor vouchers for 
cited illustrations consists of the SCD cultigen, and most of these specimens 
are of the pilose-morph (see preceding section). Therefore, overturning the 
choice of LINN 783.4 is not supported. 


Correct name of the widespread SCD cultigen 
To rectify the common practice of misapplying the name Lantana camara to 
the widespread SCD cultigen and to implement Méndez’s recommendation that 
“the system established by Sanders around L. camara should be reinterpreted,” 
an attempt has been made to locate, to examine and, when needed, to choose 
nomenclatural types of all other published names in sect. Lantana (App. B). 
The results demonstrate that other names, also, have been misapplied by a num- 
ber of specialists, including Schauer, Moldenke, Sanders, and Méndez. 

With regard to the abaxial leaf indument, the SCD cultigen is essentially 


392 BRIT.ORG/SIDA 22(1) 


outside the range of variation of Lantana camara but within the range of the 
Strigose Group (Fig. 4, 12). Thus, it would be imprudent to submerge this widely 
naturalized, aggressive species of hybrid origin as an infraspecific taxon of L. 
camara. Rather, the SCD cultigen more likely has a complex parentage involving 
L. nivea Vent. L. scabrida Sol. in Aiton, and L. splendens of the Strigose Group, as 
well as L. camara, and possibly other species (noteably, L. hirsuta of the Setose 
Group). Morphologically, it retains only the leaf shape and adaxial surface dull- 
ness of L. camara. Given the protracted history of collection and publication of 
new taxa, it is surprising that none of the available names can be verified as ap- 
plying to this widespread cultigen. Apparently, this is due to there being so many 
published names and to the confused state of the taxonomy of sect. Lantana. 

Through uncritical analysis of vague descriptions, one could argue that a 
few of the names that lack known types could be applied to the cultigen. How- 
ever, taxonomic stability is more important than unsubstantiated priority. As- 
signing a neotype toa name whose concept can never be known with certainty 
in order to honor early taxonomists, who never understood the complexity of 
these plants, would be imprudent. Such names were never taken up and have 
no precedence of usage in the horticultural and ecological communities that 
taxonomists attempt to serve. 

As a species of hybrid origin, of which that origin remains obscure, the 
SCD cultigen must be treated as a species nova. To provide taxonomic and bib- 
liographic continuity, thus maximizing stability, the new epithet proposed here 
is based on the root “-camara.” To indicate its hybrid nature and complex hy- 
pothesized parentage involving several species of the Strigose Group, it is given 
the prefix “strigo-"to produce Lantana strigocamara. At least two infraspecific 
epithets, and perhaps others, could serve as basionyms. However, epithet prior- 
ity is mandatory only within rank, and I chose a new species name. Lantana 
camara L. var. rubella Moldenke (1949) is not widely known, and applying it to 
the whole of the species would broaden Moldenke’s concept considerably. Lan- 
tana flava Medik. f. sandersii Méndez (2002) is only four years old and has not 
been taken up outside Méndez’s own work. Raising it to species level would be 
tantamount to honoring myself. 


— 


Lantana strigocamara R.W. Sanders, sp. nov. (Fig. 4). Type U.S.A. Floripa: Dade Co. 
near intersection of Montgomery St. and Old Cutler Rd, Montgomery Botanical Center, dis- 
turbed edge of rock pineland, 23 Sep 1981, Sanders 1450 (HOLOTYPE.: FTG! [Dig. photo! see 

Fairchild Tropical Botanic Garden 1999}, IsoTYPE: NY!; Drawing, Sanders 1987, fig. 9). 


Credita hybrid le Lant d L.etL.nivea Vent. et L.scabrida Sol. in Aiton et L. splendenti 
Medik. et L. hirsuta M.} ns& ti mixta simulat speciem. Differta 1 ‘a laminis 
paces euis str dees s, ichomatibus remotis non nisi nervis mediis secondariis tertiariisque 


geniculatis; a L. nivea et L. scabrida et L. splendenti laminis 


magis rugosis, ee abruptius contracti ypInnInery ibus, oe adn hebetibt bus ma nitidis; a L. 
at Ns 


nivea trichomatibus | vel aureis; a 


o x t 


SANDERS, TAAUNVIVIT UF LANTANA SECT. LANTANA 393 


amnelieaue | 


Strict to lax, ae erect, iid or ene shrub to 3 m tall (climbing 
much higher in some areas of the paleotropics); twigs hirtellous, angled, with 
or without prickles. Petioles 5-10 mm long; blades cordate to ovate, 3-10 x 2-6 
cm, mostly 1-L7 times longer than wide, the base abruptly narrowed onto peti- 
ole, the apex acuminate, moderately rugose between higher orders of veins, dull 
above; hairs of adaxial laminar surface 0.3-0.7(-1.2) mm, scattered, in the form 
of antrorse strigae, often from pustulate bases; hairs of abaxial laminar surface 
usually 0.5 mm or less, stout, in the form of attenuately conical, geniculately 
antrorse strigae, but not closely appressed to surface, not deciduous, usually 
restricted to and scattered on midrib, secondary and tertiary veins (in some 
populations small erect hairs 0.1-0.2 mm long weakly developed on smallest 
veins and along crevice under larger veins); abaxial sessile glands green or not 
visible with 10 lens; teeth 15-25 per side, obtuse to short-acuminate, not revo- 
lute. Inflorescences 2-3 cm in diameter; peduncles 2-10 cm long; receptacle 
fistulose; bracts mostly 3-8 mm long, subulate, long-triangular or narrowly 
lanceolate (single outer series sometimes longer and narrowly spathulate or 
rarely subfoliaceous), apically attenuate, puberulent or hirtellous abaxially only, 
deciduous in fruit. Calyx 3 mm long, membranous; corolla yellow, orange-red, 
white, or pink to deep reddish purple and often mixed with orange or crimson, 
the throat often yellow, the tube 7-12 mm, slightly curved, the limb 6-10 mm 
across. Drupe ca. 5 mm across, blue-black; endocarp obovoid, inflated. 


Paratypes: BURMA: Chantaburi, Makam forest, 26 Aug 1966, Larsen et al. 1841 (AAU, LL!). CHINA. 
Kwangsi: Kweilin, cult., 1979, Wan & Chow ates L!). INDIA. Lucknow, cult., 28 Oct 1974, Shivarajan 
85750 (LL!). PAPUA NEW GUINEA. East Sepik . Wewak, 31 Aug 1968, Krauss 1287 (LL!). SRI 
LANKA. Sabaragamuwa Prov.: Kegalle: Klnahatene 12 Feb 1974, Moldenke et al., 28331 (LLLUSI). 
UGANDA. Paraa, 29 Jul 1972, Moldenke & D ke 26073(LL!). USA. Florida. Dade Co.: Homestead, 
18 Jul 1966, Smith FI. 6 (BRI, LL!). New York. nate Co.: Yonkers, cult., 23 Sep 1950, Moldenke 
21133 (LL!) 


Hypothesis of the history of the cultivated Lantana camara complex 

Prior to the Linnaean period, the following species (citations in App. B) were 
already in cultivation in Europe, as deduced from the Sherard specimens at OXF, 
the Sloane and Clifford Herbaria at BM, and the Linnaean Herb. at LINN and S: 


Pane a (Fig. 1, West Indies and Mexi h outh America) 


Pilose group Lan 
Lantana. horrida (Fig 8, aes aa id, Mexico to eons 
Strigose group Lantana nived (Fig. 7, southern Brazil to ae 
Lantana scabrida (Fig. 2, West Indies, Mex 
Lantana splendens (Fig. 6, Bahamas) 


Setose group — Lantana hirsuta (Fig. 3, Mexico) 


All these species, except L. nivea, are characterized by capitula single in the 
leaf axils with stout peduncles and with corollas opening yellow and changing 


394 BRIT.ORG/SIDA 22(1) 


Fic. 6. Lantana splendens, epitype (Herb, Sherard 1269, OXF), a. infl f i b. adaxial leaf 


surface.c. abaxial leaf surface. Scale bars:a = 1 cm;b &c= 1 mm. Photogray pplied by yf 


to darker yellow, orange, or red-orange, and are aculeate or not. To my knowl- 
edge, Lantana nived is exceptional among the natural species in having capitula 
commonly paired in the leaf axils with filiform peduncles and with corollas 
opening white and aging bluish, rose or purple, or opening pink and aging pur- 
plish. The plants are usually aculeate with recurved prickles, sometimes fiercely 
so. Therefore the only source in the cultivated complex for capitula opening 
yellow or cream and changing to purplish reds or orange plus purple would be 
hybridization between L. nivea and one or more of the other species. Parentage 
involving L. nived is also consistent with stoutly recurved prickles in some of 
the cultivated hybrids. Apparently by backcrossing L. camara into such hybrids, 
plants with the mixed-colored capitula were developed within the range of 
variation of L. camara with respect to other characters (Fig. 9). Such plants | 
consider to be the subspecies combination L. camara subsp. aculeata. lam rec- 
ognizing subspecies here and elsewhere in the complex for the following rea- 
sons: |) the rank varietas in Lantana sect. Lantana has been overused for culti- 
vars, and the varietal names are largely misapplied; 2) there may be yet 
undiscovered variety epithets that would take priority and further destabilize 
the nomenclature; and 3) | recognize the subspecies rank for geographically 
widespread infraspecies that comprise varieties, and further research will likely 
reveal localized geographic races that may warrant recognition at the rank of 
varietas. According to the ICBN (Greuter et al. 2000), this hybrid taxon cannot 
be treated as a nothosubspecies, but rather Art H3.3 would require treating it at 
the nothospecies level. However, it does not behave as a species distinct from L. 
camara, and it is the major representative of the species where it is naturalized, 
thus, simulating a widespread natural subspecies. 


Lantana camara L. subsp. aculeata (L.) R.W. Sanders, comb. & stat. nov. (Fig. 9). 
BASIONYM: Lantana aculeata L., Sp. Pl. 627. 1753. Lectotype designated by Méndez & Cafferty 
(2001, see App. B). 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 395 


“s 
ry 
»* 
“* 
a 


Fic. 7. Lantana nivea, ef tative | infl itt jJuncle, and inset of 


ve {Cioht 
t 


t 
eres fy £ 


, C),a. stem I 
inflorescence. b f surf c. abaxial leaf Scale bars:a =1cm;b&c=1mm. 


Likewise, backcrosses of L. nivea into the original mixed-color hybrids produced 
multicolored plants within the range or extending the range of variation of L. 
nivea with respect to other characters. Such plants I consider to be the subspe- 
cies combination L. nivea subsp. mutabilis. It is not treated as a nothotaxon by 
similar reasoning. 


Lantana nivea Vent. subsp. mutabilis (WJ. Hook.) R.W. Sanders, comb. & stat. 
NOV. BASIONYM: Lantana nived Vent. var. mutabilis WJ. Hook., Bot. Mag. 5: pl. 3110. 1831. LEc- 
TOTYPE (here designated): icon in WJ. Hook., Bot. Mag. 5: pl. 3110. 1831. 

Lantana nivea subsp. mutabilis may have been an important genetic vector for 

the origin of L. strigocamara. Even so, this taxon is not to be identified with L. 

strigocamara regardless of Bailey’s 1900) and Moldenke’s (1980b) having com- 

bined this taxon under L. camara and applying it to L. strigocamara (Moldenke 

in sched.) (see L.amethystina and L. mutabilis Lippold ex Otto & A. Dietr, App. B.). 

The array of species names, dating from the Linnaean and early post-Lin- 
naean period, that apply to hybrids (App. B, Fig. 12) shows the extent to which 
horticultural hybridization was complicating the systematics of this group. 

Lantana camara subsp. aculeata was hybridized at least with L. horrida (pro- 

ducing L. x mutabilis C. E. Weigel), with L. hirsuta (L. X mista L., Fig. 10), and 

with L. strigocamara or some undetermined species or species hybrids from 
the Strigose Group (L. x purpurea Hornem.). Lantana horrida was crossed with 


396 BRIT.ORG/SIDA 22(1) 


K 


Se 
. 


Fic. 8. Lantana horrida.a. portion of lectotype (Humboldt & Bonpland 4149, P-HBK, photograph provided by the Herbier 
National Paris [P], used with permission). b—c. representative specimen (Laughlin 1535, LL). b. adaxial leaf surface. c. 
abaxial leaf surface. Scale bars: a = 1cm;b&c=1mm 


some of the species or hybrids of the Strigose Group to produce L. xflava L. 
(Fig. Ll). This latter hybrid appears to have mixed with backcrosses of L. nivea 
into L.camara subsp. aculeata (L. x antidotalis Schumach. & Thonn.). Further- 
more, within the Strigose Group, L. nivea was hybridized probably with L. 
scabrida or L. splendens to yield L. x multiflora Otto & A. Dietr. 

At about this time, these various hybrid plants were being introduced and 
naturalized throughout the tropics, especially as colonists settled in the West 
Indies, India, South Africa, Indonesia, Australia, and Pacific Islands. In some 
cases, the hybrids underwent polyploidy resulting in cultivated and escaped 
plants that grew aggressively and genetically swamped the characters of other 
cultivated and escaped plants with which they crossed. Thus, over the next two 
centuries, the diversity of the wild escaped plants was reduced to a few aggres- 


SANDERS LANTANA SECT, LANTANA 397 


a 


Fic. 9. Lantana camara subsp. aculeata, epitype (BM-SL 98: 143 bottom center ).a. whole specimen. b. adaxial leaf sur- 
face. c. abaxial leaf surface. Scale bars = 1 mm 


sive cultigens and the recombinant variation expressed in hybrids between 
cultigens and in those between cultigens and indigenous species. 

Apparently having originated along with the other early cultigens, Lan- 
tana strigocamara became one of the most widespread and aggressive. Clearly 
it was already escaped and crossing with native L. urticoides Hayek of Texas 
and L. depressa of Florida by the 1820s (see L.x rubra Berland. and L.xfloridana 
Raf., App. B). Lantana strigocamara probably originated as a diploid, because 
diploid cultivars, usually given names such as ‘Dwarf Yellow, ‘Dwarf Pink, etc., 
are still available (Sanders 2001). However, post-origin allopolyploidy certainly 
has contributed to its aggressive growth and success in the wild. In introduced 
areas, it appears to have steadily increased in numbers, and by the early 1900s it 
came to dominate the occurrences among pantropical naturalized plants. 

Howard (1969) pointed out that during the later half of the nineteenth cen- 
tury, lantanas lost popularity among horticulturalists. Most of the early hybrid 
cultivars apparently went extinct, except where escaped and naturalized in the 
absence of competition with Lantana strigocamara. However in the 1950s, lan- 
tana cultivars, developed primarily from a base of L. strigocamara, began to re- 


398 BRIT.ORG/SIDA 22(1) 


a 


Pe pe rT +A € ct +t 


J 
I last £ 
GHOMAIOUS I€dl 


Fic. 10. Lantana x mista, epitype (Herb. Sherard 1272, OXF). a. whol ‘ 
hap nas Javinl laaf curt Scale bars:a = 1¢m;b,c 


2. 
h 


b. inflorescence. c 


gain popularity, especially in seasonally dry subtropical areas where they can 
be used as drought tolerant borders. In particular, Monrovia Nursery in Califor- 
nia crossed L. strigocamara with the very drought tolerant, persistently bloom- 
ing L. depressa var. depressa of Florida to produce the now widely planted 
Callowiana Hybrids (Sanders 2001). 

The Callowiana Hybrids apparently originated after autotetraploidy was 
induced in Lantana depressa var. depressa. The resulting tetraploid L. depressa 
would hybridize with tetraploid L. strigocamara to produce hybrids with an 
even balance of characters from both parents. This hybrid was then hybrid- 
ized with various cultivars of L. strigocamara to generate an array of cultivars 
with a wide variety of floral color combinations, growth habits, and ploidal 
levels from triploid to hexaploid (Sanders 2001). The Callowiana Hybrids, thus, 
comprise a spectrum of variation completely connecting L. strigocamara and 
L. depressa var. depressa. This confusing variation led Moldenke (1975b) to name 


— 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 399 


va } a : 
\ : 
4 : Ney 
aa {\S 
. Ve ee 
oy 4 * 


Fic. 11. Lantana x flava, epitype (Herb. Sherard 1275, OXF). a. infl 1 rey ive | b. adaxial leaf 
surface. ¢. abaxial leaf surface. Scale bars:a = 1¢cm;b &c=1mm. 


one of these hybrids, probably Lantana cv. ‘Cream Carpet, as L. bahamensis f. 
albiflora. 


CONCLUSION AND SUMMARY 


The convoluted horticultural and natural history of Lantana sect. Lantana has 
resulted in highly complex systematic relationships of both cultivated plants 
and plants growing in the wild. Initial focus on cultivated forms during the Lin- 
naean period resulted in an early proliferation of names for closely related cul- 
tivated plants and hybrids. These names clouded the taxonomy of naturally 
occurring species as plant exploration penetrated remote areas away from hu- 
man population centers in colonial tropical America. Horticultural selection 
developed aggressively growing allopolyploid cultigen species and subspecies 
that became naturalized, often as pernicious weeds. 

Even though Lantana strigocamara (i.e., the SCD cultigen), one of the most 
widespread and aggressive of these cultigens, is characterized predominantly 
by technical features of the Strigose Group, it became confused with L. camara 


400 BRIT.ORG/SIDA 22(1) 


S 
& 
i) 
@ 
S 
— 
Q 
wn 
= 
a 
Oo 
wn 
@ 
Or @ 
’ HH » =e | PE RRO eas 2. sone ata a _ 
‘ Mutabilis ; 2 
. uv a cc 
Sean auc - : =s 
‘antidotalis | 
+ 
Fic. 12.Diag howing relationships I speci ' { hybrids. Dashed ovals indicate hybrids tt 
are extinct or rarely seen, either Itivated plant turalized pes. Solid indicate | likely 
+ kK a aA Cah 4 1; £ |} , | nh L J J 1 
bs J ede tec A 


dod d £ } } H | . £. 4s 4 


Tl i re +h £+h + +h 


setose-morph plants on the left. The ‘‘is omitted from hybrid names. For full names see Appendix B. 


of the Pilose Group. Otto & Dietrich (1842) applied to one color variant of Lan- 
tana strigocamara the name L. mutabilis Lippold ex Otto & Dietr, but this isan 
illegitimate later homonym of L. mutabilis C. E. Weigel, a name published for 
the distinct hybrid combination L. camara subsp. aculeata X L. horrida (both 
of the Pilose Group). Sanders (1987a, b, c, 1989a, b, 2001) and Méndez (2002) 
both saw the need to treat the widespread SCD cultigen as a distinct species. 
Unfortunately, Sanders misapplied the name L. camara to the SCD cultigen and 
the name L. urticifolia to L. camara. Méndez correctly applied the name L. 
camara but misapplied L.xflava to the cultigen. Thus, the SCD cultigen is newly 
named and described here as L. strigocamara. 


SANDERS LANTANA SECT. LANTANA 401 


As popularity of the original cultivars waned, later horticultural selection 
used a restricted set of aggressive polyploid hybrids, especially Lantana strigo- 
camara, as breeding stock; thus, this species of hybrid origin and its hybrids 
now strongly dominate the horticultural trade. Furthermore, where it has been 
introduced into tropical America, it competes or also hybridizes with indig- 
enous taxa to produce a spectrum of variation challenging taxonomic research, 
which may be resolvable only with cytological and molecular techniques. 


APPENDIX A 


Use of name Lantana camara L. in taxonomic and horticultural literature in 
which trichomes are sufficiently described, in historical sequence. 


Pilose-morph only 


Meoikus (1775) “folia...minus rugosa, mollioraque foliis Lantanae mistae” 

SpRENGLE (1825) “subtus albido-villosis” 

SCHAUER, (1847) “subtus pallidis pagina vel reti saltem brevi-villoso subcanescentibus” 

ScHAUER, (1851) diagnosis = “subtus pallidis pagina v. reti saltem brevivilloso 
subcanescentibus" and description ="Foliorum indumentum variabile, magis pilosum 
et (subtus imprimis) magis tomentosum, magis minusve d tamen semper breve 
et in facie foliorum scaberrimum,e setulis antrorsis callo insidentibus confectum, neque 


hirsutum’ 

Troncoso (1965) “densamente villoso-pubescentes abajo” Note:Lantana camara does not 
naturally occur in Argentina. Thus, this description applies to escaped cultivars or to 
plants of Lantana horrida (incl. L. tiliifolia), L. micrantha, and L. riedeliana, which are na- 
tive but submerged by Troncoso into L. camara. 


Pilose- and strigose-morphs both included 


Voss (1896) “oberseits scharf, unterseits blasser odor auf der Unterflache oder ihrem 
Adernetz kurz-grauweif-zottig.” [inclusion of wide range of hair morphs clear from 
synonymy, included varieties, and use of “scharf” to modify “zottig.”] 

BaiLey (1900) “pubescent beneath” [inclusion of wide range of hair morphs clear from 
synonymy and included varieties.] 

STANDLEY (1924) “variously pubescent or occasionally glabrate beneath 

HUTCHINSON & Datziet (1931) “shortly pubescent below” 

Baitey (1951) “pubescent beneath” [inclusion of wide range of hair morphs clear from 
synonymy and included varieties.] 

LEON & Atain (1957) “el envés pubescente” [term equivocal, could be applied to both morphs] 

Maceribe (1960) “often canescently short-villous beneath” [presumes less pubescent mor- 
phs also included] 

GiBson (1970) “variously pubescent beneath (densely viscid-tomentose to strigose or rarely 
glabrate)" 

Lopez-Patacios (1977) varying from “glabresentes’ to “esparcido-pilosos o estrigulosos en 
el envés" to “corto-pubescentes por ambas caras” 


" 


402 BRIT.ORG/SIDA 22(1) 


NasH & Nee (1984) “el envés densamente viscido-tomentoso a estrigoso o hirsuto, 
raremente glabro" 

Huxtey (1992) “pubesc. above and beneath” [inclusion of wide range of hair morphs clear 
from synonymy and included varieties.] 

CULLEN ET AL. (2000) “downy or almost hairless underneath” 

Strigose-morph only (or also mixed) 

Orto & Dietrich (1841) diagnosis = “subtus hirtulis” and description = “oberhalb durch 
kleine Héckerchen scharf, etwas kurzhaarig, unterhalb ebenfalls [emphasis mine] 
zerstreut kurzhaarig, und etwas heller grun oder beinahe weiflich [glaucous?]” Note: 
They described Lantana moritziana, which is conspecific with true L. camara, as “Die 
Blatter... kurz-weichhaarig, nur wenig scharf und ziemlich weich anzufahlen.” Further- 
more, their description of L.camara is almost identical to theirs for L.crenulata, which 
Schauer (1847) described as “subtus pallidis, pubescenti-scabris.” 

Pearson (1912) “scabrid-pubescent above and on the veins beneath" 

Mo oenke (1970a) “short-pubescent, mostly on the venation beneath 

Lone & Laketa (1971) “strigose beneath" 

Apams (1972)"Lower surface of leaves thinly pubescent or almost glabrous" (as opposed 
to“with numerous short hairs’) 

Mo toenke (1973b) “usually only sparsely pilosulous or strigillose beneath (mostly only on 
the venation) with canescent or brownish trichomes, sometimes glabrescent” [as op- 
posed to “densely puberulent or short-pubescent’] 

Corrett & Corrett (1982) short pubescent, mostly on the venation beneath” 

MOLDeNke & MOLDENKE (1983) “Usually only sparsely pilosulous or strigillose beneath (mostly 
only on the venation) with canescent or brownish trichomes, sometimes glabrescent” 

Jansen-Jacoss (1988) “strongly or sparsely hirsute-strigose below, especially on the veins” 

Waaner €T AL. (1990) “lower surface sparsely short-pilose or strigillose, sometimes glabrate” 

Verocourt (1992) “+ pubescent or glabrescent beneath but scabrid on raised venation" 

Lioaier (1994) “algo pelositas en el envés, a veces glabrescentes" 

Poot (2001) “envés con tricomas estrigosos pequenos limitados a los nervios” [My work, 
as well as LOpez-Palacios (1977) cited.] 


7] 


APPENDIX B 


Typification and history of application of names in Lantana sect. Lantana. 
Unless otherwise noted, herbaria given for authors and collectors in TL-2 
(Stafleu @ Cowan 1976-1988) were consulted for original material. An exhaus- 
tive search for isotypes was not attempted. Unless cited otherwise, handwrit- 
ing comparisons based on Burdet (1979). A name is considered to be misap- 
plied if the cited author or annotator clearly misunderstood the concept of that 
species and applied it toa species in sucha way to exclude the type. A misappli- 
cation is inferred for publication of an infraspecific combination in which the 
infraspecific taxon actually belongs toa second species that excludes the type 
of the species under which the combination is made. The hybrid formulae for 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 403 


nothospecies and species of hybrid origin are presumed parentages based on 
character combinations and oe 


1753—Lantana camara L., Sp. P|. 6 
Lectotype (Moldenke & Moldenke 1 i —Cult, probably Hort. Uppsala, Herb. Linnaeus 783.4 (LINN!) 
ig. ] 
Name misapplied to 
L. xantidotali (Verdcourt 1992) 
L. xflava (Moldenke 1980b) 
L. horrida ae 1962, 1982c; es 2002) 
L. X mista (Bailey 1900; Moldenke 1942) 
L. nivea (Bailey 1900; Moldenke eve 1980b) 
L. scabrida eae ide 1973b; Adams 1971) 
L. splendens (Molden 
L. strigocamara (Otto . te 1841; Moldenke 1940, 1949, 1970a, 1973b, in sched.; Long & Lakela 
1971; Correll & Correll 1982; Sanders 1987a, b,c, 1989a, b, 2001, in sched; Verdcourt 1992; Liogier 
1994) 


Taxonomic disposition.—L. camara (Pilose Group) 
Discussion.—Linnaeus' concept of L.camara was developed from an array of garden and horticul- 
turally selected plants. His reference in the protologue, in Sp. Pl. (ed. 2) 1763, and in Syst. Nat. (ed. 12 
1767 to synonyms that include the phrases “floribus miniatus,"“flore variegata,” and “flore variabile’ 
indicates that garden hybridization had already introduced genes of at least L. nivea into the mate- 
rial he studied. However, the Moldenkes’ lectotypification associated the name with an element 
characteristic of native Jamaican populations that lack the wider Genene ae and have only 
flowers opening yellow and changing to darker yellow, or at most, to r Indeed, Proctor 
18266 (NY!) and Yuncker 17043 (NY!), native material from Jamaica, are close orticg to LINN 783.4. 

Otto and Dietrich (1841) appear to have been the first to misapply L.camara to L. strigocamara 
They described L. camara as having leaf blades (my translation) “basally somewhat rounded or 
almost cordate and having surfaces above with small, rough, sharp, somewhat short hairs, and 
likewise below scattered short-haired.’They restricted the application to material of L. strigocamara 
with flowers opening yellow and aging orange (see also L. crocea [1804], L. moritziana [1841], L. 
mutabilis Lippold ex Otto & A. Dietr. [1842], and L. variegata [1842] below) 

rs (1987¢) explicitly excluded the type of L.camara by describing the species as having, 

“Hairs of abaxial laminar surfaces sparse, restricted mostly to midrib, secondary, and tertiary veins, 
stout, tapering-conical, geniculate toward base with distal 2/3 held parallel to lamina or vein surface.” 


1753—Lantana aculeata L., Sp. P|.627. 
Lectotype (Méndez & Cafferty 2001).— icon in Plukenet, Phytographia t. 233, f.5. 1692. 
Epitype (here designated).—Herb. Sloan 98:143 bottom center specimen (voucher of Plukenet, 
ee 233, aa BM-SL! (Fig. 9) 
Name misapplied to 

a camara Greats 1775; Méndez 2002) 

a X L. strigocamara (Méndez 2002) 

hy horde Boch ees ann 1858; Méndez 2002) 

L. nivea (Méndez 

L. strigocamara ne a 
Taxonomic beatae on.—L. camara (as L.camara subsp. aculeata) 
Discussio omy knowledge ae (1847) was the first author to place L. aculeata in synonymy 
under ie camara, which has equal priority, and his choice is followed. 

sndez and Cafferty chose Plukenet's figure as the lectotype. The voucher of the lectotype 

(i.e, epitype) is more or less within the range of variation of L. camara with regard to indument. 


= 


—t 
— 


404 BRIT.ORG/SIDA 22(1) 


Although the abaxial surface we are ot wee is typical praia material, this condition is 
found in occasional native collec nnaeus y developed the concept of L. 
aculeata to accommodate ae? that therwise like L.camara but differed only by the devel- 
opment of prickles. As with L. camara, a. proteleeds indicates both plants with yellow flowers 
changing only to dark yellow ate those with flowers opening yellowish changing to red or pur- 
plish.In particular, Plukenet’s ame includes “floribus miniatis.” Not only the floral color varia- 
tion (especially the late ee Salah pigments from L.nivea), but also the presence of prickles 
and sparse abaxial foliar hairs both point to hybridization influencing the wider variability in L. 
aculeata. Thus, my concept of the taxon is that of an infraspecific rank to account for all the culti- 
vated and escaped plants of hybrid as (regardless of floral color) falling within or slightly ex- 
sae the range of variation of L. camara. 
éndez described L. aculeata as mon surface with hairs relatively abundant, usually on all 

veins fa eres areoli), robust, conical, all, most, or at least a significant portion on the midrib, sec- 
ondary or tertiary veins geniculate toward base and with distal 2/3 held parellel to vein surface, all 
those that grow on lower veins shorter and erect.”To me, this describes hybrids between L. camara 
and L. strigocamara. Therefore, his application of L.aculeata to L.camara x | sn 
conform to the voucher of the lectotype, pointing out the need for epitypification. Aaa L. 
aculeata f. rubella (Moldenke) I.E. Méndez is to be identified with L. strigocamara, L. aculeata f.nivea 
(Vent.) LE. Méndez with L. nivea, and L. aculeata f. parvifolia (Moldenke) I.E. Méndez hae L.camara 

This taxon apparently was the main vector for infusing L. camara genes into the Strigose 
Group to produce L. strigocamara. It is not now widely available in the roticltutal trade but ap- 
pears to be rather widely naturalized in the paleotropics, especially Austra 


1767—Lantana mista L., Syst. Nat.ed. 12, 2:417 
Lectoryee (Méndez & Cafferty 2001).—icon in pienie Hort. Eltham. t. 56, f.64. 1732. 
Epitype (here designated).—Herb. Sherard 1272. (voucher for Dillenius, Hort. Eltham. t. 56, f.64) OXF! 
(Fig. 1 
Name misapplied to 

L. hirsuta os hauer 1847, 1851 t. 42) 

L. horrida (Méndez 2002) 
eee disposition.—L. x mista (L. hirsuta x L.camara subsp. aculeata) 
Discussion.—Reliance on the Dillenius figure led Méndez (2002) to associate this name with the 
wild species, L. horrida, which occurs in the Greater Antilles. However, when the epitype (i.e, voucher 
of the figure) is examined with reference to critical trichome characters, it is shown to be a hybrid 
between . hirsuta and L. camara subsp. aculeata. It is not typical of L. horrida. 


2s 


hx} ri oa et 


This hybrid combination is mostly of historical interest, as it is not now widely available in the 
horticultural trade and may have served as an agent for further hybridization and introduction of 
wider genetic variation into the cultivated complex 


1768—Lantana pio Mill, Gard. Dict. ed. 8, Lantana 5. 
LecTorre (Sanders 1989a).—Mexico. Edo. Veracruz: Veracruz, Houstoun s.n., Herb. Sloane 6:84. (BM-SL 
photo!, mounted epHotes NY!, UCI). 

cme misapplied to 

L.can cara Gueldenke in sched.; Adams 1971; _ 1987b, c, 1989a, b; Liogier 1994) 

L. horrida (Adams 1971, Sanders 1987b, 1989 
Taxonomic disposition.—L. camara (see 1753) 
Discussion.—Mark Spencer la Typification Project, pers. comm.) verified that Herb. Sloane 
6:84, collected by Houstoun at Veracruz, Mexico, is the only specimen bearing Miller's polynomial 
and is the only known original oa of L. urticifolia. Nash & Nee (1984) cited the collection as 
“type” but questioned its existence at BM. Based on a mounted photograph at UC of the specimen 
at BM, Sanders (1989a) cited the specimen as “holotype” which is to be corrected to “lectotype.” 


oo 
= 


SANDERS LANTANA SECT. LANTANA 405 


Because Sanders (papers cited pov) had lest L.camara so as to exclude its type, he 
applied the name L. urticifolia t ies name to all members of L.camara.|In his concept 
of L. urticifolia, he also included the species a horrida, ne regarded as distinct. 


1775—Lantana flava Medik., Hist. & Commentat. Acad. Elect. Sci. ae a 3.Phys. 225, 

Lectotype (here designated).—icon in Dillenius, Hort. Eltham. t. 57, f.6 
Epityee (here designated).—Herb. Sherard 1275 (voucher for aa on ENAaAy: t.57, f.66). OXF! 
(Fig. 11) 
Name misapplied to: 

L. camara an 1847; Koch & Fintelmann 1858) 

L. scabrida (Méndez 2002) 

L. strigocamara nee 2002) 
Taxonomic disposition.—L. flava (L.horrida X L. spp. Strigose Group) 
Discussion.—There is no indication on the Sherard specimen that Medikus saw it, and his descrip- 
tion closely parallels Dillenius: Because the protologue cites only Dillenius' name in synonymy, the 
Dillenius plate must be the type 

The protologue clearly ascedbes the lower leaf surface as “pallidiora, tomentoso-pilosa.” | 


bearing the mixture of long and short appressed hairs on the foliar upper surface and mixture of 
antrorse strigae with softer, filiform hairs on the lower surface, oe epitype (. e,, uch of the lec- 
totype) agrees with the protologue and confirms that the rrida and one 


or more species of the Strigose Group. Therefore, it is not clear why Méndez (2002) ae to apply 
this name he sate ta of L. ee He cited neither the plate nor its voucher. 

ti mostly of historical interest, as it is not now widely available in the 
ovticuturel trade and may ne served for further hybridization and introduction of wider ge- 
netic variation into the cultivated complex. 


1775—Lantana splendens Medik., Hist.& Commentat. Acad. Elect. Sci. Theod.-Palat. 3. Phys. 226. 

Lectoryee (here designated).—icon in Dillenius, Hort. Eltham. t. 57, f.67. 1732 

Epitype (here designated).—Herb. Sherard 1269 (voucher for Dillenius, Hort. Eltham. t. 57, f. 67), left 

hand stem. OXF! (Fig. 6) 

Name male p Rte to: 
ie 


pe 


foldenke & Moldenke 1983) 

Tepenonie dis pou on.—L. splendens (Strigose Group) 

Discussion.—The lectotypification by Moldenke & Moldenke ([Dillenius s.n., OXF] 1983) is rejected 
as both imprecise and incorrect. The protologue cites as original material only the Dillenius name 
and plate as a definite synonym and a Plukenet name and plate as a questioned synonym. Not only 
does Medikus’ description closely parallel Dillenius, but the voucher is not annotated by Medikus. 
Therefore, there is no evidence that Medikus studied the specimen, and the Dillenius plate is cho- 
sen as lectotype. Both the protologue and the epitype (i.e., the voucher of the aa rnatch 
native material from the Bahama Archipelago, the provenance given by Dilleni 


1775—Lantana sanguinea Medik,, Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 3. Phys. 229. 
Type.—Unknown. 
Name misapplied to: 

L. camara or L. X mista (Schauer 1847) 
Taxonomic disposition.—L. camara subsp. aculeata (see 1753) or L. X mista (see 1767) 
Discussion.—Medikus cited no synonyms, illustrations, or specimens that could serve as type. His 
own herbarium is unknown. His description of trichomes is scant, making clear application of the 
name difficult. He did relate it most closely to L. aculeata L.; thus, the assumption made here is that 
it is a color variant of that taxon or L. mista L., for which he gave a similar description, except for 
ultimate color of the corollas. 


406 BRIT.ORG/SIDA 22(1) 


1776—Lantana mutabilis C.£. Weigel, Physiogr. Sallsk. Hand. 1 

Lectorype (here designated).—Cult., Greifsweld Bot. Gard. ("H. ae '), Pyl s.n., Aug. 1774 UE (dig. 
photo!)). 

Taxonomic disposition.—L. x mutabilis (L. camara aes aculeata ° x L.horrida) 

Discussion.—The protologue indicates that the n the Greifsweld Botanic Garden 
was the basis of the new species. The 1774 collection a Pyli is the a erieinal material found at JE 
and was annotated by Weigel U. Mueller, JE, per ly, Wei ited Camara melissae 
folio, on aaa Dill. (Hort. Eltham.65, t.56, f.65),in synonymy.The Het ee ee illustration (Herb. 
Sherard 1274, OXF!) also is a hybrid between L. camara subsp. aculeata and L. horrida. 


=f 
— 


1789—Lantana scabrida Sol.in Aiton, Hort. Kew, ed. 1, 2:352. 

Lectoryee (here designated) —Cult. Royal Bot. Gard. Kew,"Hort. Kew 1777," (Solander s.n.?) (BM). (Fig. 2) 
Taxonomic disposition.—L. scabrida Clears Grou p) 

Discussion.—The specimen at BM apparently is the only original material. It is annotated “Hort Kew 
1777" in an unknown hand, possibly Solander’s, and “Lantana scabrida Ait. Hort. Kew ii 352!" possibly 
by J. Britton (M. Spencer, BM, pers. comm.). The only material cited in the protologue is “Nat. of the 
West Indies, Mr. Gilbert Alexander, Introd. 1774." 


1789—Lantana melissifolia Sol. in Aiton, Hort. Kew, ed. 1, 2:352. nom. illeg. (superfluous) 

Lectotype (here designated).—icon in Dillenius, Hort. Eltham. t.57,f.66.] 

Epitype (here designated).—Herb. Sherard 1275 (Voucher for Dillenius, re Eltham. t.57, f.66). OXF! 
Ieee rai —l. Xflava (see 1775) 

Discussion.—Solander cited Lantana flava Medik. in synonymy and cited the Dillenius plate. Evi- 
dence is oe that he studied the voucher in the Sherard Herbarium. 


1796—Lantana mutabilis Salisb., Prodr. Stirp. Chap. Allerton. 107. nom. illeg. (later homonym of L. 
x mutabilis C.E. Weigel) 

Type.—None selected. 

Taxonomic ne tion.—L. camara subsp. aculeata (see 1753) 

Discussion.—Because this is a later homonym ake xmut gall is C.E. Weigel, it presents no problems. 
Based on the protologue, Salisbury app ial with heads that change colors 
from yellow to purple. He cited only L. camara Linnaeus in Sp. Pl. (ed. 2) 1763 in synonymy; so, it is 
not clear whether he was ae a substitute name or separating the material with multicol- 
ored heads as anew spec 


1804—Lantana crocea Jacq,, P|. Hort. Schoenbr. 4:t.473. 
Lectotyee (here designated).—icon in Jacq., Hort. Schoenb. 4:t. 473. 
Name misapplied to: 
L. bahamensis (Schauer 1847; oo & Fintelmann 1858) 
L. camara (Otto & Dietrich 184 
ibrida (Sck 1847 se . Fintelmann 1858) 
L. sple ndens cone 1847) 
Taxonomic disposition. —L. camara (see 1753 
Discussion.—No original material other than the plate has been found. The plate is a good match 
for many wild collected specimens from Jamaica. These differ from the type of L. camara only by 
details of leaf shape. However, herbarium samples suggests (unpubl. observ.) that, among Jamai- 
can Penn: leaf-shape variation is not correlated with Baa ae or ecology. 
Bec e Otto and Dietrich (1841, 1842) applie d L.camara to L. strigocamara (see above), they 
applied i crocea to L. camara, restricting its use to ee with heads changing from yellow to 
orange and stems with prickles (see L. moritziana, 1841 


w 


1804—Lantana nivea Vent., Jard. Malmaison t.8. 
Lectoryee (Here designated).—icon in Vent., Jard. Malmaison t.8. 


— 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 407 


Epirype (here designated).—Cult., Paris “ex h. Paris, mis. Siebke,” Siebke s.n, C! (Fig. 7) 
Taxonomic disposition.—L. nivea (Strigose Grou 
Discussion.—Ventenant provided a good description, excluding some details of leaf indument. Al- 
though the garden at Malmaison received material from gardens in the East Indies, it is clear that 
the description and plate apply to native material from southeastern Brazil, as noted by Schauer 
(1847). None of Ventenant’s ofginal matetlal sea in Paris was located in B-WILLD, C, G, or P 
(no response from WU), cultivated in Paris was annotated as L.nivea, but 
apparently not in Ventenant’s hand, sae it not to be original material. Because subsequent 
authors have misunderstood this species as evidenced by the several new names described for 
more recent wild collections of this species, the specimen in C is designated as to support the icon. 
It clearly matches the protologue and was at Paris about the time of Ventenant such that the 
annotator should have been familiar with Ventenant’s concept 


ae 


1815—Lantana purpurea Hornem., Hort. Bot. Hafn. 2:583. 
Lectotype (here designated). —Cult. Hort. Reg. Bot. Hafn."1814" (Cl). 
Name ee ed to: 

ra (Schauer 1847; Koch & Fintelmann 1858; Bailey 1900) 
eee eee —L. X purpurea (L.camara subsp. aculeata x L. spp. Strigose Group) 
Discussion.—The lectotype is in a type folder marked as “specimina originalia” (also labeled as "IDC 
microfiche 111"'*). The verso is annotated “Lantana purpurea Horne. 1814,” of ean in 
Hornemann’s hand, and stamped “Hb. Schum..” A second specimen (IDC microfiche 1 a is anno- 
tated “Lantana purpurea h.h.” in a different hand and stamped “Hb. Liebm.” Both s mens are 
characterized by a mixture of erect and geniculate trichomes on the abaxial eae purplish 
corollas suggest that a purple-flowered cultivar of L. camara subsp. aculeata was crossed with a 
cultivar from The Strigose Group 


1817—Lantana hispida Kunth in HBK, Nov. Gen. Sp. 2:260. 
Lectoryee (Nash & Nee 1984).—Mexico. Edo. Veracruz:"juxta Xalapam,” Humboldt & Bonpland s.n. (P- 
K (dig. photo!, Macbride Neg. oe Fl, BRIT!)). 
Name misapplied to 
L. hirsuta (Moldenke 1947, 1973b) 
L. hirta Grah., sp. sect. Callioreas (Schauer 1848; Koch & Fintelmann 1858; Standley 1924; 
rea 1963, roe 1970) 
L. velutind Mart. & Gal. sp. sect. Callioreas (Moldenke 1982c) 
Taxonomic ee —L. horrida (see next) 
Discussion.—Schauer’s misapplication of the name to L. hirta Graham in section Calleoreas has re- 
sulted in ee confusion. Furthermore, the epithet is easily confused with L. hirsuta M.Martens 
& Galeotti, a distinct species. Therefore, in combining L. hispida with L. horrida as conspecific,| choose 
the latter in accord with ICBN Article 11.5 (Greuter et al. 2000). 


1817— pants ae Kunth in HBK., Nov. Gen. Sp. 2:2 
Lectotype (Nash 84).—Mexico. Distrito Federal:“monte eChealpecees urbem Mexici,.” 
Humboldt &Bonpand4i 49 (P-HBK [dig. photo!]). (Fig. 8a) 
pals misapplied to 
rubra Cahaue 47) 

L pee ee 1942, nae . 1978) 

L. hirsuta XL. camara (Moldenke ) 
Taxonomic Deen as horrida ae Group) 
Discussion. —Berlandier 2310, annotated by Berlandier as Lantana rubra (L. urticoides X L. 
strigocamara), a syntype of L. horrida var. parviflora Schauer, and the epitype of L. x rubra (see be- 
low) was mistakenly included in L. horrida by Schauer (1847). As a result, Moldenke misapplied the 
name L. horrida to L. urticoides until correcting himself (Moldenke 1978). 


408 BRIT.ORG/SIDA 22(1) 


1827—Lantana antidotalis Schumach. S Thon in poeta eek ou PI.276. 

Lectotype Junghans 1962.p.94,as to collec h plicat Ghana, Thonning 
25 (C [IDC microfiche ident. no. 64 | *"]!; soLectoryee: ClIDC. no.64'°"]!). 

ae disposition.—L. xantidotalis (L.nivea X L.camara * L. x flava?) 

Discussion.—The duplicate most in eae with the protologue (leaf blades 2-3 inches long) is 

chosen as the lectotype. Though given the same collection number, these two specimens may 

actually represent two different populations as judged from slight morphological differences. As- 

signing hybrid status to this naturalized plant is based on the variable leaf shape and size (narrowly 

ovate-oblong or narrowly triangular;length 1.7 to 2.7 X width), subsetiform trichomes on the up- 

per and lower leaf surfaces,and the mixture of erect and gemicuiste hairs on the lower leaf surfaces. 

The parentage probably includes L. nivea or L.splendens from the Strig nd L.camara and 

L. X flava from the Pilose Group. 


Y 


829—Lantana albopurpurea Desf, abl. Ecole Bot., ed 3. 393. 
Lectotype (here designated).—Cult. Hort. Paris.,"H. p.” labeled “Herbarium Webbiana ex Herb. 
Desfontaines” (FI [dig. photo!]). 
Taxonomic disposition.—L. X mista (see 1767) 
Discussion.—The specimen at Fl is clearly original material, as the annotation is in Desfontaines’ 
hand (C.Nepi, pers.comm.). None other is known to me. The leaf indument suggests this specimen 
has the same parentage as does L. X mista. 


1829—Lantana suaveolens Desf, Tabl. Ecole Bot., ed 3.393.nom. illeg. (non L. suaveolens Spreng. = 
L. angustifolia Mill, 
Type.—Not investigated. 

Taxonomic disposition.—probably L. camara or its hybrid (see 1753) 

Discussion.—The description of prickly stems, ovate leaves, lanuginose bracts, and yellow flowers 
suggests this is an element of L.camara 


te 


32—Lantana rubra Berland. in Teran & Berland., Mem. Comis. Limites 15. 
LECTOTYPE . designated).—icon in Berlandier in Ohlendorf et al., transl. Journey Mex., t.5 (top, 
facing p.410).1 
Epitype (here designated). —Berlandier 2310=880, Mexico. Tamaulipas: Matamoros, GH! (isoepitype: NY, 
none found at G under L. rubra). 
Taxonomic disposition. ie xrubra (L.urticoides X L. strigocamara) 
Discussion.—The only original material known to me includes the recently published plate, which 
Berlandier either executed or supervised, and three sheets (Berlandier 2114=697 [GH!], 2310=880 
([GH!, NY!]), which consist of hybrids. Although the protologue suggests L. urticoides (see 1906 be- 
low), the only native species in the region of the eastern Texas-Mexico boundary, all the original 
material lacks the rotund, large-toothed leaf blades and persistant fruiting bracts, characteristic of 
L. urticoides. Instead, they are intermediate between L. urticoides and L. strigocamara, which obvi- 


rial that actually belongs to L. urticoides should be found, such a pein could be easily 
overturned. However, stability of the application of L. rubra and rete Uke urticoides as the 
correct name of the wild species is best served by accepting the hie as i 

the two poten omen: aa A me, a 14= er" is more ane ee hybrids, espe- 
cially in the ho 1.5mm long. Berlandier 23 10=880 
has pioeneeth ve supeersctent bracts more like the L. urticoides parent, but the indument shows 
it to be a hybrid. It is chosen as epitype because it is annotated “Lantana rubra B,” apparently in 
Berlandier's hand, is a syntype of L. urticoides var. parvifolia Schauer, and is represented by dupli- 
cates in major herbaria. 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 409 


This hybrid combination is the typical cultivated plant in Texas and the Gulf Coast Plain of the 
Southeastern United States. It may have been propagated for the trade due to the cold hardiness 
ae from the Texas parent, L. urticoides 
x Kewensis Jackson 1895) cites’ agian rubra Perr.,”but this is a mistake as Perrottet (1825) 
ca referred to Latania rubra Jacq. (Arecaceae), the next entry in his list after Lantana. 


1832—Lantana tiliifolia Cham., Linnaea 7:1 
Lectoryee (Santos Silva 2001).—Brazil: Bahia. Si ae (B-WILLD 11502 [dig. photo! Berlin Dahlem. 


005). 

Name misapplied to: 
L. strigocamara or its hybrids (Moldenke in sched) 

Taxonomic disposition.—L. horrida (see 1817) 
Discussion.—The protologue lists three syntypes—the Sieber collection from Bahia, Brazil, Sellow 
collections (“southern Brazil...copious numbers from many locations and times,” BR [dig. photo]; 
NY!, fragment NY!),and Lhotsky s.n (n.v,). | disagree with the lectotypification by Santos Silva (2001). 
In the digital photograph of B-WILLD 11502 provided by B, the specimen appears to have mostly 
glandular polis on sy stems ang petigl>: Ne Sellow specimen at BR bears mostly non-glandular 
hairs. Chamisso's diagnosis ly _ hirtis,"although his description iy ansietion) 
indicates that the age stems have Teac ie often but not always mixed in.” Later, Poeppig 
segregated the strongly glandular plants as L. glutinosa (see 1842 below). Modern usage follows 
Poeppig in restricting L. tiliifolia to the non- or weakly glandular specimens. However, because | 
treat both L. tiliifolia and L. glutinosa as synonyms of L. horrida, further research is needed to deter- 
mine whether a proposal to conserve a different type is neede 


1832—Lantana floridana Raf., Atl. J. 148. 
Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, Pl, WIS, or WS; no response from LE, NAP or W. 
Taxonomic disposition.—L. x floridana (L. depressa x L. strigocamara) 
Discussion.—The only native wild taxon of the upper Atlantic coast of Florida is L. depressa var. 
floridana Gone R.W. Sanders (see 1905, below). However, the protologue describes the flow- 
ers as “versicolor, yellow, orange, red, crimson on same shrub.” As discussed in Sanders (1987a), none 
of the varieties of L. depressa is characterized by the orange or red floral pigments. However, this 
does characterize hybrids between L.depressa and L. SE (the latter in Sanders 1987a as L. 
camara).In light of the hybridization sees introduce and native L. urticoides in 
Texas in the 1820s (see L. rubra 1832 above), L. strigocamara was likely also naturalized in Florida by 
that time, and Rafinesque's description refers to hybrids 


1838—Lantana antillana Raf, Sylva Tellur. 82. 

Type.—Not found in DWC, FI, G, NY, P P-DU, PH, Pl, WIS, or WS; no response from LE, NAP, or W. 
Taxonomic disposition.—L. horrida (see 1817) or L. camara (see 1753) 

Discussion.—Rafinesque noted this to be Lantana camara of authors and an Antill b, which 
he saw living. He also noted met it cles from L. fon primarily by the stem and leaves hirsute 
and the bracts ovate-lan er of these characters appears to be consistent 
with this being an element of L. ees 


1838—Lantana rosea Raf, Sylva Tellur. 83. 

Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, PI, WIS, or WS; no response from LE, NAP. or W. 
Taxonomic disposition.—probably L. reticulata Pers. or L. involucrata L. (. Sect. Callioreas) 
Discussion.—Even though Rafinesque placed this species in his subgenus Camara Raf., his subgen- 
era are neither well demarcated nor consistent. He included L. camara L.in subgenus Camara Raf. 
and the conspecific L.aculeata L.in subgenus Periana Raf. (along with L. involucrata). Lantana rosea 
is described as a native Antillean plant with small roundish tout apical and basally acute) leaves, 
rounded cinerous stems, and short, ovate bracts, and rose-p all of which appear to place 
it in sect. Callioreas. 


410 BRIT.ORG/SIDA 22(1) 


1838—Lantana incarnata Raf, Sylva Tellur. 83. 

Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, PI, WIS, or WS; no response from LE, NAP.o 

Taxonomic disposition.—probably L.nivea subsp. mutabilis (WJ. Hook.) R.W. Sanders 1804 and 

text 184 

ee on. ae incarnate,’ Rafinesque presumably meant that the heads are flesh-colored in bud, 
p ream and turn ign or fleshy pink. This is consistent with some populations of L. 

strigocamara. However, the ovate-oblong leaf blades acuminate at both apex and base rule out L. 

eal The floral color, leaf shape, and long, linear-lanceolate bracts are all consistent with L. 

vea subsp. mutabilis. 


amethystina ) 


1839—Lantana coccinea G. Don, Hort. Brit., ed. 3.245.nom. nud. 
Type.—Unknown. 
Taxonomic disposition.— 


camara subsp. aculeata? 


1841—Lantana crenulata Otto & A. Dietr,, Allg. Gartenzeitung 9:363. 
Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. 
Hort. Berlin [Otto s.n.,?] existed in B, now destroyed; Shauer [1847] indicated seeing a live specimen 
in Hort. Berlin). 
Name misapplied to: 

L. lar nei as r 1847) 
Taxonomic disposition.—L. Xantidotalis (see 1827) ar L. scabrida (see 1789 
Discussion.—The See: cites no original material, but the description strongly suggests L. 
scabrida or related species or hybrids in the Strigose Group. Identity with L. strigocamara is ex- 
cluded because the leaves are described as pointed on both ends. 


1841—Lantana pals Otto & A. Dietr.,, Allg. Gartenzeitung 9:369. 
Lectotype (Schauer 1857, as to gathering; here designated, as to specimen).—Venezuela. Caracas, 
Moritz 163 (G [dig. photol]; isotectorype: G [lacking inflor.,dig. photo!) 
Name misapplied to: 
L. camara (Otto & Dietrich, 1841, 1842) 

Taxonomic disposition.—L. camara (see 1753) 
Discussion. —The protologue seo nativity only by “Habitat in Caracas” without specifically cit- 

ng the Moritz specimen. In his treatment, Schauer (1847) cited only Moritz 163 from Caracas, Ven- 
ee but stated that he saw ce and wild-collected herbarium specimens at B. The only 
duplicates of Moritz 163 known to me are at G.The one chosen as lectotype bears two annotation 
labels besides the collection label, all apparently in Schauer’s hand. The duplicate at G lacks 
infloresences, is not annotated by Schauer, but is annotated by Moldenke as “Isotype.” Macbride 
photographed material at B (neg. no. 17478, destroyed [photo F!, BRIT!, GH!]), which he may have 
thought was type material. The specimen is not signed by either Otto & Dietrich or Schauer, and it 
bears the date’1844."Therefore it cannot be considered a type, even though Lopez-Palacios (1977 
cited it as such. 

Because Otto and Dietrich (1841, 1842) misapplied L. camara to L. strigocamara, they applied 

L.moritziana to plants of L.camara with no prickles and flower heads changing from yellow to orange. 


1841—Lantana multiflora Otto & A. Dietr, Allg. Gartenzeitung 9: oa 

Lectorype (here designated).—Cult. Hort. Berlin, Otto s.n.(B,d brid 17379 F!, BRIT!]). 
Taxonomic disposition. —L. X multiflora (probably L.nivea X L.scabrida or L. splenden 
Discussion.—Otto and Dietrich cited no specimen, but the destroyed specimen Seueehee by 
Macb oe appears to have been the only original material. In general, the protologue suggests L. 
nivea, although, the more distinct scabrosity of the leaves suggest L. scabrida or L. apIENGeN The 
floral color, described further in Otto and Dietrich (1842) as lilac (in bud?) changing to yellow, clearly 
implicates genes of both L. nivea and yellow-flowered species of the Strigose Group. 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 411 


1841—Lantana amethystina Otto & A. Dietr,, Allg. Gartenzeitung 9:370 

Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. 
Hort. Berlin [Otto s.n.,?] existed in B, now destroyed). 

Taxonomic disposition.—L. nivea (as L. nivea subsp. mutabilis, see 1804 and text) 

Discussion.—Otto and Dietrich (1842) indicated that this is a replacement name for L. nivea var. 
mutabi lis. Hooker. Apparently, Otto and Dietrich were not aware of L. mutabilis C.E.Weigel and L. 
mutabilis Salisb. but did consider that Lippold had named L. mutabilis Lippold ex Otto & A. Dietr. for 
a distinct species such that the Hooker varietal epithet was unavailable at the rank of species (see 
next entry). 


1842—Lantana mutabilis Lippold ex Otto & A. Dietr,, Allg. Gartenzeitung 10:314.nom. illeg. (later 

homonym of Lantana X mutabilis C. E. Weigel) 

Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. 

Hort. Berlin [Otto s.n.,?] existed in B destroyed; no material of Lippold found at RB [R.Campostrini 

F,, pers 

eee series —L. strigocamara (see 2006 and text) or L. strigocamara x L. camara subsp. 
ulea 


ae 


eee noted under L. camara, Otto and Dietrich (1841) appear to have applied the name 
L.camara to the phenotypes of L. strigocamara having floral heads changing from yellow to orange. 
However, material of L. strigocamara with purplish and yellowish pigments occuring together they 
apparently described under L. mutabilis Lippold ex Otto & A. Dietr.As a later homonym, this name is 
not available for L. strigocamara. 


1842—Lantana variegata Otto & A. Dietr., Allg. Gartenzeitung 10:314 

Tyee.—Unknown (no original material known; if herbarium specimen from material cultivated in. 

Hort. Berlin [Otto s.n.,?] existed in B, now destroyed). 

Taxonomic disposition.—L. camara subsp. aculeata or L. camara subsp. aculeata x L. spp. Strigose 
ro 


Discussion.—No holotype or syntypes are cited; however, the protologue suggests cultivated ma- 
terial in the range of L. camara subsp. aculeata (see 1753). 


1842—Lantana glutinosa Poepp. in Otto & A. Dietr,, Allg. Gartenzeitung 10:315. 

LECTOTYPE renee as 1977, as to gathering; here designated as to specimen).—Peru, Poeppig 
1375 (G-DC [dig. photo!]; duplicate, B?, destroyed [Macbride Neg. 34340 F!, BRIT!]; soectorype: GH! 
Taxonomic disposition.—L. horrida (see 1817) 

Discussion. —Other than eee in eae no material is cited and lectotypification is required. 
The eee photograph (Neg. 34340) is presumed to be of a specimen at B, since destroyed, 
although no herbarium eae is visible in the image. If this be true, it would have been the 
preferred lectotype. 


oot 


1842—Lantana asperata Hort. ex Vis., Orto Bot. Padova 142.nom. nud. 
Type.—Unknown. 

Taxonomic peel mes vee (see 
Discussion.—V i | | this name at Hort.Parisiense, 
without fAcading to ube it as his own.He also stated,"Non pare diversa della L. nivea Vent.” 


a 


804) 


1843—Lantana bartramii Baldwin, Reliq. Baldw. 247. 

Type.—Not designated. 

Taxonomic disposition.—L. X floridana? (see 1832) 

Discussion.—The protologue cites no material.In a search of DWC, LASCA, MANCH, NY, P-DU, PH, 
WECO and WLU, only two kee specimens were located. At NY originally in the Torrey Her- 
barium (T. Zanoni, pers.comm.) is a specimen (!) labeled “grown in Louis Lecontes’ Garden in Geor- 
gia” without collector or ie and annotated as the “Lantana camara [var] bartramii, cultivated in 


412 BRIT.ORG/SIDA 22(1) 


Florida by Dr. Baldwin."This specimen may or may not have been seen by Baldwin. At PH, there is a 
sheet (dig. photo!) from the Muhlenberg Herbarium that has mounted on it several fragmentary 
specimens from different collectors.One such fragment is labeled,”1482 Lantana Hardy native’ and 
presumably was collected by Baldwin. Further research is needed to determine if this specimen is 
original material. Both specimens are elements of |. depressa Small var. floridana (Moldenke) R.W. 
Sanders (see 1905). 

Baldwin's protologue refers to plants along the Florida seacoast as being the same that Bartram 
described, without mentioning cultivated material. Bartram traveled only in the geographic range 
of var. floridana. Further, Baldwin describes the species as attaining the height of six feet. Material of 
L. depressa var. floridana that | have seen reaches three feet but not six. This, therefore, suggests 
Baldwin was describing the same hybrid that Rafinesque did under the name Lantana x floridana. 


1844—Lantana hirsuta M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11:326. 
Hovotyee.— Mexico. Edo. Veracruz: Jalapa, Mirador, 1840, Gallioti 749 ( BR [dig. photo!]; isoryee: G [dig. 
to!]). 


pho 
Taxonomic disposition.—L. hirsuta (Setose Group) 


1846—Camara cau Benth., Bot. Voy. Sulphur 154 

Lectorype (here designated).—Herb. Linnaeus 783.4 (LINN). 

Taxonomic disposition.—L. camara (see 1753) 

Discussion.—This was a replacement name for Lantana camara when placed in Camara Adans. 
However, the correct name in Camara. is C. aculeata (L.) Kuntze (Rev. Gen. Plant. 2:503. 189 


184 ae anane amare Schauer, Linnaea 20:480. 

pel (her cited as “isotype” by Lopez-Palacios 1977).—Venezuela.’ad Caracas," Moritz 
92 (BM, n.v.; HoLoTYPE NOW destroyed: B [Macbride Neg. 17463 FI, GHI]). 

. misePPe ed to: 


LA ] 


ia 


97 2a) 
Tavonannic disposition —L. hirsuta (see 1844) 


1847—Lantana polyacantha Schauer in DC., Prodr. 11:597. 
Hototype.—Mexico, Schiede s.n. (B, destroyed, ie c 17481 [F!]). 
Taxonomic disposition.—L. horrida (see 1817) 


1847—Lantana aoe Schauer in DC,, Prodr. 11:599. 
Lectoryee (S —Brazil. Matto Grosso, Manso s.n., Mart.Fl. Bras. 1026 (M,n.v.ISOLECTOTYPES: 
BM, fe ) (dig. ee c DC [dig. photo!, Macbride Neg. 7873 F!, BRIT!], G [dig. photo!, Macbride 
Neg. 24632 FI, eis K,NY!). 
Name misapplied to 

L. nivea (Moldenk Ke ae 

L. viscosa (Moldenke 19 
Taxonomic disposition. —L. en (Strigose Group) 
Discussion.—The are two paratypes known to me. One is Poeppig 1485 (Peru) at G. It apparently is 
annotated by Schauer (without his “!"). He cited Poeppig 1405 which is probably a typographical 
error for"1485."The other is Martius s.n. (Brazil. Rio ely at M (n.v.). 


1847—Lantana robusta Schauer in DC., Prodr. 

Lectorype (Santos Silva 2001).—Brazil. Rio de cea i ae do Correia,” 1821, Poh! 40-5955 (W, nv). 
Taxonomic disposition.—L. robusta (Setose group) 

Discussion.—There is a Macbride photograph ae 17485 [F!, BRIT!, GH!, LL! NY!J) of Pohl 782 at B, 
now destroyed, that has been distributed as a photograph of a type. The specimen in the photo- 
graph should be disregarded as original material. This specimen neither appears to be annotated 
by Schauer nor bears sufficient inflorescence material to match the protologue. Furthermore, the 
paratype, Raben 509 (NY [fragment]! and BR [photo at LL!]), and Poh/ 182 are not the same species. 


SANDERS LANTANA SECT. LANTANA 413 


Pohl 182 may belong to L. horrida or L. horrida X L. robusta. 


1847—Lantana riedeliana Schauer in DC., Prodr. 11:601. 
Hototype.—Brazil, Riedel s.n. (LE, n.v.). 
Name misapplied to 

L. sp. sect. ee loreas (Moldenke 1970b) 
ce ep osion: —L.riedeliana or L.robus 
Discussion.—The only original material cited is a sigeate at LE, which did not respond to inquiries. 
There is a photograph in LL (perhaps by Moldenke) of Glazier 1657 that is annotated as L. riedeliana 
by Moldenke, as well as someone other than Schauer. This specimen appears to fit the description 
of L. riedeliana and suggests that L. riedeliana is conspecific with L. robusta, being a glabrescent, 
more narrow-leaved variant. 


1847—Lantana pohliana Schauer in DC., Prodr. 11:601. 

Type (Lectotype not designated).—Brazil, Pohl s.n. (W?, n.v.; duplicate: B, destroyed [Macbride Neg. 
17480 F!, BRIT!, GH!]). 

Taxonomic disposition.—L. pohliana or L. viscosa? (Setose Group?) 

Discussion.—Schauer cited no collection number or herbarium. Pohl material should be at W, but 
no response to inquires there was received. Further investigation is neede 


1847—Lantana viscosa Pohl ex Schauer in DC., Prodr. 11:601. 

Lectotype (Santos Silva 2001).—Brazil. Goids:"Ad Concei¢do, prope Trahiras,” Poh! 1876 & 2680“D. n° 
181." (left-hand specimen) (W, n.v.;isoLectoryee: B, destroyed [Macbride Neg. 17492 FI, BRIT!]) 
Taxonomic disposition.—L. viscosa (Setose Group) 


— 


1847—Lantana multicolor Lem., Fl. Serres Jard. Eur. 3:239 


O 
L. camara subsp. aculeata or its hybrid (Koch & Fintelmann 1858) 

Taxonomic disposition.—L. camara subsp. aculeata, L. x mista, or L. sp. sect. Callioreas? 
Discussion.—Seeds of this were sent to Brussels from Mexico, possibly of cultivated origin. Lemaire 
distinguished it from other cultivated lantanas by its unusually large leaves and by the rose and 

rimson flowering heads. Although Koch & Fintelmann (1858) recognized the species, they indi- 
cated that it may be a form of L. camara or L. Xmista. However, the protologue also describes the 
heads as involucrate with four large bracts and with the floral bracts cordate-rotund. These charac- 
ters would place the plants in section Callioreas 


1857—Lantana hybrida Neubert, Deutsch. Mag. Garten- Blumenk. 10:98. nom. illeg. (nom. subnud.) 
Lectoryee (here designated). —icon in Neubert, Deutsch.Gart. Mag. 10:t. facing p. 1 
Name diese ed to 

Le ara subsp. euleane or L. ee (Everett 1980) 

ee disposition.—L. X flava (see 1775) or L. x mista (see 1767) 
Discussion.—No specimens were found at ‘oe or STU to serve as lectotype or epitype. The de- 
Se pHenN: is very brief, indicating only the color of the flower heads. The plate shows nothing dis- 
tinctive except for spreading hairs on the twigs, petioles,and peduncles, suggesting that the plants 
may be elements of L. flava or L. X mista. 


1858—Lantana es K. Koch & Fintelmann, Wochenschr. Gartnerei Pflanzenk. 1 :322.nomm.illeg., 
pro syn. sub L.cro 

Tyee—Unknown. 

ees ic disposition.—L. camara (see 1753) 


1863—Lantana triplinervia Turcz,, Bull. Soc. Imp. Naturalistes Moscou 36:205. 
Hovotyee.—Java, Goering 225 (KW?, n.v.). 


414 BRIT.ORG/SIDA 22(1) 


Name misapplied to 

L. nivea (Moldente 1974, 1977) 

L. nivea X L. spp. Pilose Group (Moldenke 1974, in sched; Sanders in sched.) 
Taxonomic disposition.—L. nivea (see 1804) 
Discussion.—Turczaninow eae material that was cultivated or escaped from cultivation. The 
protologue strongly suggests L. nivea by the leaves long petiolate, ovate-lanceolate, triplinerved, 
apically acuminate, and minutely scabrous on the veins below. 


1863—Lantana bahiensis Turcz., Bull. Soc. Imp. Naturalistes Moscou 36:206. 
Hototype.—Brazil. Bahia, Sa/zmann s.n. (KW?, n.v.). 
Name misapplied to: 
L. nivea X L. spp. Setose meee mani in sched.) 
Taxonomic disposition. —L. sp. sect. Callior 
Discussion.—The protologue are ne species in section Callioreas, which is corroborated by 
the description of bracts ovate, briefly acuminate, and 5 ed. 


1904—Lantana micrantha Briq., Annuaire Conserv. Jard. Bot. Geneve 7-8:299, 
Ho.oryre. ve taatley Asuncion, Balansa 1039 (G). 
Name misapplied to 
L. horrida (Moldenke 1948, 1975d) 
L. sp. sect. Call ie 1981c) 
Taxonomic disposition.—L. micrantha (Pilose Group) 
1905—Lantana depressa mall, Bull. New York Bot. Gard. 3:436. 
HoLoryPe. ee. en a Co.:pinelands between Coconut Grove and Cutler, Small & Carter 747 
(NY; isotype: F!). 
axonomic disposition.—L. depressa (Strigose Group) 
Discussion,—See Sanders (1987a) 


1905—Lantana bahamensis Britton, Bull. New York Bot. Gard. 3:450. 
Hotoryee—Bahamas. New Providence: Ft. Montague, Britton & Brace 174 (NY!;\sotyees: F!, US! 


= 


Name misapplie die 

L. camara (Moldenke 1975a; Correll & Correll 1982 (pro parte)) 

L. depressa (Moldenke 1975c) 

L. depressa X L. strigocamara Callowiana Hybrids (Moldenke, 1975b) 

L. splendens (Moldenke in sched; Correll & Correll 1982 (pro parte); Sanders in sched.) 
Taxonomic disposition.—L. bahamensis (sp. hybrid origin between L. splendens and L.camar 
Discussion.—Apparently Lantana camara, occurring in the southern Bahamas, began to hybridize 
introgressively into L. splendens, centered in the central Bahamas, shortly after settlement by Euro- 

As a result, the native plants in the Bahamas are variable. The type of L. bahamensis shows 
the nitid surface and scattered geniculate hairs on the leaf undersurface venation typical of L. 
splendens, as well as the soft, erect hairs persisting along the undersurface veins, apparently ex- 
pressing genes of L. camara. The bracts vary from oblong-lanceolate as in L. splendens to obovate- 
elliptic as in L.camara 


1905—Lantana ovatifolia Britton, Bull. New York Bot. Gard. 4:123. 
Hotorype.— Bahamas. Grand Bahama: Eight Mile Rocks, Britton & Millspaugh 2450 (NY1; soryee: FI). 
Name misapplied to: 

L. depressa (Long 1970; Long & Lakela 1971; oes 1982) 


L. strigocamara X L. depressa (Long & Lake 71) =L. xfloridana Raf. 
Taxonomic disposition. eo ovatifolia isos ni 
Discussion.—See Sanders 1987a. 


1906—Lantana glandulosissima Hayek, Repert Spec. Nov. Regni Veg. 2:161 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 415 


Hotorvee.—Mexico. Jalisco: Tequila, Pringle 4431 (W, n.v.isotypes: BR, F!, MO!, NY!). 
Name Sa ere 
Wet 


4 (KAS -] 


L. 
Taxonomic disposition.—L. camara (see 1753) 


1906—Lantana cummingiana Hayek, Repert.Spec. Nov. Regni Veg. 2:161 
Hototyee.—Chile. Cumming 1065 (W, n.v.) 
Taxonomic disposition.—L. horrida = 1817) or L. hirsuta (see 1844) 


1906—Lantana urticoides Hayek, Repert.Spec. Nov. Regni Veg. 2:162. 
Tyee—Not selected, to be lectotypified in a subsequent paper. (Syntypes: USA: Texas. Lincheimer 
Exsic. No. 503, which includes two different Lindheimer numbered collections, F!, GH!, MO!, UCI, W, 
n.v.; Texas: Comal “ Matthes 19, W,n.v,). 
Name misapplied 
L. horrida ae 1981b) 
Taxonomic disposition. —L. urticoides (Setose Group) 
Discussion.—See L. rubra 1832 above 
1907—Lantana foetida Rusby, Bull. NewYork Bot. Gard. 4:431. 
Hototyee.— Bolivia, Bang 2034=469 (NY [n.v., dig. photo not available]; isoryees: US!, GH!). 
Taxonomic disposition.—L. horrida (see 1817) 


1909—Lantana weberbaueri Hayek in Urb., Bot. Jahrb. Syst. 42:166. 
Hotoryee.—Peru. Junin: Tarma: Palca, Weberbauer 2017 (GB?, n.v., W?, n.v., or B? [destroyed, Macbride 
Neg. 17493, LL). 

Taxonomic disposition.—L. weberbaueri or L. viscosa (see 1847) or its hybrid? (Setose group) 
Discussion.—This is a rare, poorly known species. Hayek's opinion was that it is near L. viscosa and L. 
robusta in the Setose Group of section Lantana 


1910—Lantana arida Britton, Bull. Torrey Bot. Club 37:357. 
Hototyee.—Jamaica, Fort Henderson, Britton & Hollick 1824 (NY!). 
Name misapplied to: 
L. camara (Moldenke, 1982a) 
L. horrida (Moldenke, 1982a: Méndez, 2002) 
Taxonomic disposition.—L. horrida (see ) 
Discussion.—This is a small-leaved variant of Antillean L. horrida, perhaps as a growth response to 
drought conditions. 
912—Lantana subcordata Urb. Symb. Antill. 7:351. 
Lectoryee (here designated)—Dominican Rep.near Santiago, Schomburgh 5 (K! [Hotoryee:B, destroyed]; 


CO 
~~ 


ISOLECTOTYPE: P). 
Taxonomic disposition.—L. subcordata (Pilose Group) 
Discussion.—See Sanders 1989a 


1922—Lantana pulchra Larranaga, Escritos D. A. Larranaga 1:406.nom. dub. 


Type.—Destr 
Taxonomic ae —Cannot be determined. 
Discussion.—Written in the style of the Linnaeans, the protologue contains only 14 words besides 


the ee and provides no features to distinguish it from other published species. As both the 
specimens and Larranaga’s drawings are destroyed (E. A. Paz, MVFQ, pers. comm.), the name must 
be treated as a nomen dubium 

1934—Lantana tenuifolia Rusby, Phytologia 1:74. 

Hotoryee.—Bolivia. Bopi River, Rusby 653 (NY [n.v., dig. photo not available). 

Taxonomic disposition.—L. cujabensis? (see 1847) 


416 BRIT.ORG/SIDA 22(1) 


1940—Lantana insularis Moldenke, Caribbean Forester 2:16. 

Hovotyet.—Jamaica. trail from St. Helens Gap to Latimer River, Maxon & Killip 912 (NY!;\sorype: US! 
Taxonomic disposition.—L. insularis (Setose Group) 

1940—Lantana eee lceeas Ssleae et 2:17. ee as L. “leonardorum” 
Hototype.—Haiti. near Jean Rabel, | 82 (NY!; isotypes: Al, GH! US!). 
eines ion.—L. heats (Pilose pee 

Discussion.—See Sanders 198 


= 


1940—Lantana scorta Moldenke, Publ. Carnegie Inst. Washington 522:161. 
Hototype.—Mexico. Queretaro: near San Juan de Rio, Rose et al. 9520 (NY!). 
Name misapplied to: 

L. horrida (Moldenke in sched.) 

L.camarax L. horridaX L. hirsuta (Moldenke 1940) 
Taxonomic disposition.—L. hirsuta (see 1844) 


1940—Lantana notha Moldenke, Phytologia 1:422. 

Hotoryee.—Mexico. Sinaloa: Fuerte, Rose et al. 13573 (NY1). 

Taxonomic disposition.—L. notha? (section ? 

Discussion.—Although Moldenke (in sched.) applied this name to plants in section Lantana, the 
type appears to be in section Callioreas, but the critical character to distinguish sections, mature 
fruits, is lacking. 

1941—Lantana scandens Moldenke, Phytologia 2:18. 

Hototype.—Mexico. Michoacan: Coalcoman: Villa Victoria, Pto. de Aire, Hinton 12315 (NY! isorypes: FI, 
LL!, MO! NY!, US). 

Taxonomic disposition.—L. scandens or L. scabrida (see 1789 Strigose Group) 


1941—Lantana brittonii Moldenke, Phytologia 2:52. 

Hotoryee.— Jamaica. Tweedside, below Moody’s Gap, Harris & Britton 10541 (NY!:isoryee: US). 

Taxonomic disposition.—L. scabrida (see 1789) 

1946—Lantana minasensis MICIEELING) Byoiegia 2:1 38. 

HoLoryee. pea Minas ‘| ¢ B Mexia 4448a (NY [dig. photo!]; 
TEX 


ISoTYPES: M 
Name ea to: 
L. nivea (Moldenke 1946) 
L. nivea X L. spp. Pilose Group (Moldenke 1966, 197 2a, 1973a, in sched.: Sanders in sched.) 
Taxonomic disposition.—L. nivea (see 1804) 
1962—Lantana kingii Moldenke, Phytologia 8:161 
orype.—Mexico, Oaxaca: 2 km south of Niltepec, King 1775 (TEX! isotype: US!) 
Taxonomic disposition. —L. kingii or L. scabrida (see 1789, Strigose Group) 
1979—Lantana morii Moldenke, Phytologia 41.449. 
Hotoryee.—Brazil. Bahia: Mun. Camaca: Sao Joao do Panelinha, Santos & Mattos 3304 (LLI:isorype: NY! 
Taxonomic disposition.—L. nivea (see 1804) 


= 


1987—Lantana hodgei R.W. Sanders, J. Arnold Arbor. 68:343. 

Hotoryer.—Dominica. Fresh Water Lake, elfin forest, Fosberg 48269 (USI; isoryees: F!, GHI, K, MO!, NY!). 
Taxonomic disposition.—L. hodgei (Strigose Group) 

2006—Lantana strigocamara R.W. Sanders, in this publication. 

Hotorype.— USA. Florida: Dade Co., Sanders 1450 (FTGI; isotype: NY!). (Fig. 4) 

Taxonomic disposition.—L. strigocamara (L. spp. Strigose and Setose Groups  L. camara subsp. 
aculeata) 


— 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 417 


ACKNOWLEDGMENTS 


The Botanical Research Institute of Texas supported this research with institu- 
tional facilities and resources. The Queensland Department of Natural Re- 
sources and Mines provided a travel grant to support the author while in Eu- 
rope to visit BM, K, and OXF. Michael Day and Isidro Méndez encouraged the 
author to complete this research after a hiatus of several years. Logistic and 
library support provided by Charlie Jarvis is especially appreciated. Curators 
of the cited herbaria are thanked for loans, digital images, and historical infor- 
mation. Guy Nesom, John McNeill, Sandy Atkins, and Charlie Jarvis provided 
helpful discussion and comments. 


REFERENCES 


Aoams, C.D. 1972. Flowering plants of Jamaica. Univ. West Indies, Mona, Jamaica. 1972. 

BaiLey Hortorium. 1976. Hortus third:a concise dictionary of plants cultivated in the United 
States and Canada. MacMillan, New York. 

Baivey, LH. 1900. Cyclopedia of American horticulture. MacMillan, New York. 

Baivey, L.H. 1951.Manual of cultivated plants. MacMillan, New York. 

Berin-DAHLEM. 2005. Digital specimen images at the Herbarium Berolinense. April 15,2005. 
http://ww2.bgbm.fu-berlin.de/herbarium/ 

Briquet, J. Verbenaceae. 1895 ("1897"). In: A. Engler & K. Prantl, Nat. Pflanzenfam. IV. 3a: 
132-182. 

Buroet, H.M. 1979. Auxilium ad botanicorum graphicem. Conserv. & Jardin Bot., Geneva. 

Corrett, D.S.and H.B. Corrett. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz. 

CULLEN, J. eT AL., (eds.). 2000. The European garden flora. Cambridge Univ., Cambridge. 

Day, M.D., C J.Witey, J PLavrorp, and M.P. Zatuck!.2003. Lantana: Current Management Status 
and Future Prospects. A.C.LA.R. Monograph 102. Australian Centre for International 
Agricultural Research, Canberra. 

Everett, T.H. 1980. Lantana. |n:T.H Everett, ed. The New York Botanical Garden illustrated 
encyclopedia of horticulture, 10 vol.. Garland, New York. 6:1927-1929. 

FAIRCHILD TROPICAL BOTANIC GARDEN. 1999. The virtual herbarium. http://www.virtualherbarium.org/ 
5 Jan 2006. 

Fernanoes, R.B. 1988. Notes sur les Verbenaceae. VII. Remarques sur quelques especes 
africaines du genre Lantana L. Bol. Soc. Brot.61:125-214. 

Gisson, D.N. 1970. Verbenaceae. In: PC. Standley and L.O.Williams, eds. Flora of Guatemala. 
Fieldiana, Bot. 24, pt. IX(1, 2):167-236. 

GreuTer, W. eT aL. (eds.). 2000. International code of botanical nomenclature (Saint Louis 
Code). Reg. Veg. 138. Koeltz Scient. Books, K6nigstein. 

Hooker, W.J. 1831. Lantana nivea var. mutabilis. Bot. Mag. 5:pl. 3110. 

Howard, R.A. 1969.A checklist of cultivar names used in the genus Lantana. Arnoidia 29: 
73-109. 

Howarb, R.A. 1970.Lantana camara—A prize and a peril. Amer. Hort. Mag. 47:31-36. 


418 BRIT.ORG/SIDA 22(1) 


HuTCHINSON, J. and J.M. Datziet. 1931. Flora of West Tropical Africa. London. 

Huxtey, A.et at. (eds.). 1992. The new Royal Horticultural Society dictionary of gardening. 
MacMillan, London 

Jackson, B.D. 1895. Index Kewensis. 2 vol. Clarendon, Oxford. 

Jarri, SM.H.and A. GHaroor. 1974.Verbenaceae. Fl.West Pakistan 77:1—40. 

Janson-Jacoss, M.J. 1988. Verbenaceae. In: A.R. A. Gorts-Van Rijn, ed., Fl. Guianas. A. Phan- 
erogams 4(148):3-86. 

Jarvis, C.E. 1992. Seventy-two proposals for the conservation of types of selected Lin- 
naean generic names, the report of Subcommittee 3C on the lectotypification of Lin- 
naean generic names. Taxon 41:552-583. 

Jarvis, C.E., E.R. Barric, D.M. ALLAN, and J.L. Reveat. 1993. A list of Linnaean generic names and 
their types. 100 pp. Konigstein. 

JUNGHANS, J. 1961-1962. Thonning's and Isert’s collections from “Danish Guinea” (Ghana) in 
West Tropical Africa. Bot. Tidsskr. 58:310-355, 59:82-199. 

KHosHoo, T.N. and C. Manat. 1967. Versatile reproduction in Lantana Camara. Curr. Sci. 
36:201-203. 

Kocu, K. and G.A. FinteLMANN. 1858. Die Camaren Brasilien’s und Westindien’s (Lantana 
Camara und die Verwandten). Wochenschr. Gartnerei Pflanzenk. 41:321-325. 

Kuntze, O. 1891.Revisio generum plantarum. Pars Il. Leipzig. 

Leon, H.and H. Atain. 1957. Flora de Cuba. Vol. 4. Fernandez, Havana. 

Linnaeus, C. 1737. Hortus Cliffortianus. Amsterdam. 

Linnaeus, C. 1753. Species plantarum. Stockholm. 

Liocier, A.H. 1994. La flora de la Espanola. VI. San Pedro de Macoris, Santo Domingo, Dom. 


Rep. 

Lone, R.W. 1970. Additions and nomenclatural changes in the flora of Southern Florida |. 
Rhodora 72:17-46. 

Lone, R.W. and O.Laketa. 1971.A flora of tropical Florida. Univ. Miami, Coral Gables, Florida. 

Lopez-Patacios, S. 1974. Cambios de estado, reduccidn a sinonimia y registros nuevos de 
algunas Verbenaceae de Venezuela. Revista Fac. Farm. Univ. Andes 14:21-23. 

Lopez-Patacios, S. 1977.Lantana. |In:S.L6pez-Palacios. Flora of Venezuela—Verbenaceae. Univ 
Los Andes, Mérida (non series Fl. Venezuela: Caracas). Pp. 327-416. 

Maceribe, J.F. 1960. Verbenaceae In: Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser. 13, pt.5, 
no.2:609-7 20. 

McNeut, J., E.A. Ovett, LL. Consaut, and D.S. Katz. 1987. American Code and later 
lectotypifications of Linnaean generic names dating from 1753: A case study of dis- 
crepancies. Taxon 36:350-401. 

Meoikus.F.C.1775.Historia et Commentationes academiae electorialis, theodoro-palatinae, 
Vol. 3 Physicum. Mannheim 


Menoez, LE. 1992. Nueva tribu de Verbenaceae y clasificacién de los géneros cubanos. 
Fontqueria 33:1-6. 

Menpez, I.E. 1993. La tribu Lantaneae Briq. (Verbenaceae) en Cuba. Fontqueria. 36: 
245-251. 


SANDERS, TAXONOMY OF LANTANA 419 


Menoez, I.E. 2002. A taxonomic revision of Lantana sect. Lantana (Verbenaceae) in the 
Greater Antilles. Willdenowia 32:285-301. 

Menpez, I.E. and S. Carrerety. 2001. Typification of Linnaean names of taxa of Verbenaceae s. 
str.described from the Greater Antilles. Taxon 50:1137-1141. 

Mo oenke, H.N. 1940. Verbenaceae. In: C.L. Lundell, Botany of the Maya area. XVII. Publ. Car- 
negie Inst. Washington. 522:148-223. 

Motoenke, H.N. 1941. Novelties in the Eriocaulaceae and Verbenaceae. Phytologia 2:6-32. 

Mouoenke, H.N. 1942. Verbenaceae. |n:C.L.Lundell, Flora of Texas, Vol. 3. Texas Research Foun- 
dation, Renner. Pp. 13-88. (bound and reissued with other contributions, 1961) 

Mo oenke, H.N. 1946. Nomenclatural notes. Ill. Phytologia 2:129-151. 

Motoenke, H.N. 1947. Notes on new and noteworthy plants. |. Phytologia 2:213-240.— 
Additional number series, —1948.V.Phytologia 2:464-477 —1949.VIl.Phytologia 3:58- 
64.—1955.XIX.Phytologia 5:132, 133.—1962.XXX.Phytologia 8:159-164.— 1963. XXXV. 
Phytologia 9:98-100.—1966.XLIV.Phytologia 13:242,243.—1973a.LVI.Phytologia 25:220, 
221.—1974. XIX. Phytologia 28:401-404.—1975a.LXXVII. Phytologia 31:25-29.— 1975b. 
LXXIX.Phytologia 31:360.—1975c.LXXX.Phytologia 31:373.—1975d.LXXxIll.Phytologia 
32:333-336.—1976. LXXXVI. Phytologia 33:129,130.—1977. XCIX. Phytologia 36:49- 
53.—1978.CIX.Phytologia 38:498, 499.— 1981a.CXLVI.Phytologia 48:290, 291.—1981b. 
CXLIX. Phytologia 49:182.—1981c.CLIll. Phytologia 50:12-14.—1982a.CLIV.Phytologia 
50:214-215.—1982b. CLV. Phytologia 50:308-310.—1982c. CLX. Phytologia 52:129- 
132—1982d. CLXII. Phytologia 52:230-232.—1984. CLXXIll. Phytologia 55:113-115. 

Movoenke, H.N. 1954. Keys for the identification of the cultivated Verbenaceae of Califor- 
nia. J. Calif. Hort. Soc. 15:80-87. 

Mo tpenke, H.N. 1970a. Verbenaceae. In: Correll, D.S. and M.C. Johnston. Manual of the vas- 
cular plants of Texas. Renner, Texas. Pp. 1313-1342. 

Motpenke, H.N. 1970b. Further new combinations and varieties of Verbenaceae Phytologia 
19:435. 

Mo tpenke, H.N. 1972a. Novelties from Brazil, Peru, Venezuela, and Thailand. Phytologia 23: 
180, 181. 

Mo oenke, H.N. 1972b. Seven undescribed subspecific taxa in Eriocaulaceae and 
Verbenaceae. Phytologia 23:453,454. 

Mo penke, H.N. 1973b. Verbenaceae. In: R.E. Woodson, Jr.and R.W. Schery, Flora of Panama: 
Part 9. Family 168. Ann. Missouri Bot. Gard. 60:41-148. 

Mo oenke, H.N. 1980a. A sixth summary of the Verbenaceae, Avicenniaceae, Stilbaceae, 
Chloanthaceae, Symphoremaceae, Nyctanthaceae and Eriocaulaceae of the world as 
to valid taxa, geographic distribution and synonymy. Phytologia Mem. 3. 629 pp. 

Motoenke, H.N. 1980b. Reduction in taxonomic rank of some Verbenaceae and 
Eriocaulaceae. Phytologia 45:296. 

Mo oenke, H.N. and A.L.Motpbenke. 1983.Verbenaceae.In: M.D.D ke and FR. Fosberg, 
A revised handbook to the flora of Ceylon 4:196-487. 

Nash, D. L.and M. Nee. 1984. Verbenaceae. FI. Veracruz 41:1-154. 


— 


420 BRIT.ORG/SIDA 22(1) 


NaTarAjaN, A.R. and M.R. Ada. 1957. Cytotaxonomical studies in the genus Lantana. J.\In- 
dian Bot. Soc. 36:35-45. 

Orto, F.and A. Dietrich. 1841.Aufzahlung und Kultur der Lantana-Arten. Allg.Gartenzeitung 
9:362-373. 

Orto, Fand A. Dietrich. 1842. Revision der Lantana-Arten. Allg. Gartenzeitung 10:313-316. 

Pearson, H.H.W. 1912. Verbenaceae. In: W.T. Thiselton-Dyer. Flora Capensis 5:180-226. 

Perrortet, G.S. 1825. Mém Soc. Linn. Paris. 3:124. 

Poot, A. 2001. Lantana. In: W.D. Stevens et al., eds. Flora of Nicaragua (Monogr. Syst. Bot. 
Missouri Bot. Gard. 85:2511-2514. 

Sanoers, R.W. 1987a.|dentity of Lantana depressa and L. ovatifolia in Florida and the Baha- 
mas. Syst. Bot. 12:44-60. 

Sanoers, R.W. 1987b. Taxonomic significance of chromosome observations of Caribbean 
species of Lantana (Verbenaceae). Amer. J. Bot. 74:914-920. 

Sanoers, R.W. 1987c.A new species of Lantana (Verbenaceae) from Dominica, Lesser Antilles. 
J, Arnold Arbor. 68:343-348. 

Sanoers, R.W. 1989a.Lantana sect.Camara (Verbenaceae) in Hispaniola: Novelties and notes. 
Moscosoa 5:202-215. 

Sanpers, R.W. 1989b. Lantana. In: R.A. Howard. Flora of the Lesser Antilles 6:226-232. 

SAnpeks, R.W. 2001.The genera of Verbenaceae in the Southeastern United States. Havard 
Pap. Bot. 5:303-358. 

SANTOS Sitva, T.R. bos. 2001. Lectotypifications and neotypifications in Lantana and Lippia 
(Verbenaceae). Taxon. 50:1115-1118. 

ScHAueR, J.C. 1847. Verbenaceae. In: A.P. de Candolle. Prodromus 11:522-700. 

ScHAUER, J.C. 1851. Verbenaceae. In: C. Martius. Flora Brasiliensis 9:22-39. 

ScHemske, D.W. 1976. Pollinator specificity in Lantana camara and L. trifolia (Verbenaceae). 
Biotropica 8:260-264. 

Scott, L., G.C. GRAHAM, M. HANNAN-Jones; and D.K. Yeates. 1997. DNA profiling resolved the 
limited importance of flower colour in defining varieties of Lantana camara. Electro- 
phoresis 18:1560-1563. 

Spies, J.J. 1983 [publ.1984]. Hybridization potential of Lantana camara (Verbenaceae). Gar- 
cia de Orta, Sér. Bot. 6:145-150. 

Spies, J.J. 1984. A cytotaxonomic study of Lantana camara (Verbenaceae) from South Af- 
rica. S. African J. Bot. 3:231-250. 

Spies, J.J. and C.H. Stirton. 1982a.Embryo sac development in some South African cultivars 
of Lantana Camara. Bothalia 14:113-117. 

Spies, J.J. and C.H. Stirton. 1982b. Chromosome numbers of southern African plants: 1.J.S. 
African Bot. 48:21-22. 

Spies, J.J. and C.H. Stirton. 1982c. Meiotic studies of some South African cultivars of Lan- 
tana Camara (Verbenaceae). Bothalia 14:101- 111, 

SPRENGEL, C. 1825. Linnaeus’ Systema Vegetabile, ed. 16. Gottingen. 

Starceu, F.A.and R.S.Cowan. 1976-1988. Taxonomic literature, ed. 2. Vol. 1-6 (Reg. Veg. 94, 98, 
105,110,112,115,& 116) Bohn, Scheltema & Holkema, Utrecht. 


SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 421 


StanoLey, PC. 1924. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23. 

StiRTON, C.H. 1977. Some thoughts on the polyploid complex Lantana camara L. 
(Verbenaceae). In:D.P. Annecke, ed. Proceedings of the second national weeds confer- 
ence. Cape Town. Pp. 321-334 

Swarerick, J.T., B.W.Wittson, and M.A. Hannan-Jones. 1995. The biology of Australian weeds. 
25. Lantana camara L. PI. Protect. Quart. 10:81-95. 

Troncoso, N.S. 1965. Verbenaceae. In: A.L. Cabrera. Fl. Prov. Buenos Aires 5:121-140. 

Troncoso, N.S. 1974.Los géneros de Verbendceas de Sudamérica extratropical. Darwiniana 
18:295- 412. 

Verpcourt, B. 1992. Verbenaceae. In:E. Milne-Redhead and R.M. Polhill, eds..Flora of tropical 
East Africa. 155 pp. 

Voss, A. 1896. Lantana. In: A. Voss, ed..Vilmorin’s Blumengartnerei. Ed. 3, 2 vol. Berlin. Vol. 1, 
pp. 822-824. 

Waaner, W.L., D.R. Hersst, and S.H. Soumer. 1990. Verbenaceae. In: Manual of the flowering 
plants of Hawai'i 2:1315-1327. 

Wo rson, S.L. and T.W. Sotomons. 1964. Poisoning by fruit of Lantana camara. Amer. J. Dis. 
Children 107:173-176. 


422 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


STEVEN Foster and REBECCA L. JOHNSON. 2006. Desk Reference to Nature’s Medi- 
cine. (ISBN 0-7922-3666-1, hbk.). National Geographic Books, 114517" Street, 
N.W,, Washington, D.C.,, 20036-4688, U.S.A. (Orders: 202-857-7335, http:// 
shop.nationalgeographic. eee $40.00, 416 pp., 200 color photos, 150 line 
drawings, 159 maps, 7 5/8" x 4" 

This hefty book is an excellent introduction to medicinal plants for people of all ages. Co-author 

Steven Foster is a noted medicinal plant expert and photographer. So, it is no wonder that beautiful 

photographs and clear factual writing grace this collaboration between Foster and National Geo- 

graphic writer and photographer Rebecca L. Johnson. Included are 150 medicinal plants arranged 
alphabetically by common name. Plants covered include those with common household uses like 
Aloe (Aloe vera), herbal medicines like Saint John’s Wore (Hypericum perforatum), and plants that 


are sources of drugs, such as sweet asia Artemisia annua), the source of the anti-malarial 
drug artemisinin. There is even a roted toa plant used as a medicine by non-humans. The 


section on bitterleaf (Vernonia oe describes it use by chimpanzees to rid themselves of 
parasites. For each plant, the authors include brief botanical descriptions, range maps, Latin names, 
traditional and current medicinal uses, and information on cultivation and preparation. A sidebar 
in each section includes interesting tidbits about bak aa intended to add to “..a better under- 
standing of human experience with the plant or its relatives.” The useful Latin name index includes 
illustrations of each plant for visual reference. There are also essays that highlight the medicinal 
plants and ethnobotany of regions from all over the world, including Africa, North America, and the 


Middle East. Although it is not a comprehensive text book, browsing through this book is an enjoy- 
able way to ground oneself in the basics of phytomedicine and ethnobotany.—Marissa Oppel, MS, 
Herbarium Technician, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


SIDA 22(1): 422. 2006 


CRATAEBGUS SERIES PARVIFOLIAE AND ITS PUTATIVE 
ELEY BRIDS IN THE SOUTHEASTERN UNITEDSEATES 


J.B.Phipps and K. Dvorsky 


Department of Biology 
University of Western Ontario 
1154 Richmond St., North London 
Ontario N6A 5B7, CANADA 


ABSTRACT 


Dp D 


This paper deals with Crataegus series ifoli Rosaceae, subfamily Pyroideae) in the main part 
of its range, the southeastern United cues a species, C. uniflora and C. brittonii, are recognized 
together with one putative interserial hybrid, C. x vailiae. A number of other obscure entities are 


also discussed. The series and constituent species are provided with detailed descriptions, full syn- 
onymy, keys to taxa and are typified so far as possible. Line illustrations and county level distribu- 
tion maps, the latter derived from a database of over 1200 records, are also given for each species 
treated. 


Keyworbs: Crataegus, series Parvifoliae, Rosaceae ision, interserial hybrids, southeast- 


ern United States 


RESUMEN 
rE sar dae ie Parvifoliae(R subfamilia Pyroideae) en la parte princi- 
pal de su area, el sureste de I Os Bae: Unidos oS reconocen dos especies, C. kek ora re C. brittonii, 
junto con un hibrido putativo interserial, C._ vailiae. Se discuten Imbien a tidades 
poco claras. Tanto de | de | ipci letallad 


sinonimia completa, abe es de pes taxa, y se 3 tipfican en as posible para ose especie n tratada se dan 
ilustraciones y map mas de 


1200 registros. 


INTRODUCTION 


The first author was asked to prepare the Crataegus treatment for the Vascular 
Flora of the Southeastern United States only vol. 1 (Cronquist 1980) of which, 
however, was completed. In spite of the soon defunct status of this flora JBP 
continued with this Crataegus project which is now being issued as a set of 
accounts usually dealing with individual ora few series, e.g. Phipps (1988, 1998). 
The third of these is presented here and deals with Crataegus series Parvifoliae. 
This series extends well beyond the borders of the flora area, to Texas and Mexico 
westwards and New York and New Jersey northeastwards; however, it is best 
represented in the southeastern United States. Here, there is one widespread 
and abundant species, C. uniflora, which is somewhat variable, plus the local C. 
brittonii (perhaps the same as C. rhodella of western North Carolina). Several 
of the names that have been synonymised with C. uniflora nevertheless repre- 


SIDA 22(1): 423-445. 2006 


424 BRIT.ORG/SIDA 22(1) 


sent distinct nodes of variation and these will receive explicit comment here. 
Forms such as C. X vailiae and related types, however, appear to represent 
interserial hybrids with C. calpodendron (series Macracanthae) but are treated 
here for convenience. 

The primary function of this paper is to provide a taxonomic revision that 
illustrates the variation within series Parvifoliae and its putative interserial 
hybrids in a synoptic manner. Crataegus uniflora has often been illustrated 
but its more variant forms have not, as with C. brittonii, and both are illustrated 
here as well as the putative interserial hybrid C. x vailiae. None of the taxa rec- 
ognized in this paper, even C. uniflora, have ever received detailed range maps 
so far as can be determined. While the raw information to create dot maps did 
exist in our database, the scale of work necessary to provide each specimen with 
its mapping coordinates precluded such an ideal approach and instead, county 
maps, refined so as to indicate record frequency, were made. An appendix lists 
one representative specimen per county. A key to all taxa recognized here is 
also provided. The taxa recognized and their synonyms are typified so far as 
possible. However, lack of lectotypifiable material for the Ashe names involved 
has caused familiar difficulties for the typification of his names. In the cases 
encountered here, resolution through neotypification was not, however, adopted 
because of the arbitrary aspect of making type choices that would correctly 
match critically defective protologues. The situation with C. rhodella (discussed 
on p. 433) perfectly exmplifies this. Fortunately, although Ashe names consti- 
tute e the ae in series Parvifoliae, the variation encountered in the 

t d field material seen could all be accommodated within the names 
of other authors. The synonymy provided is based on names encountered in 
Palmer (1925). The paper will commence with a description of the series, fol- 
low with descriptions of the constituent taxa and close on the ¢ ive interserial 
hybrids. 

This work has been made possible by loans of over 1200 specimens from 
43 herbaria and the first author has endeavored to see and collect representa- 
tive exemplars in the field of all the taxa treated in the course of over 20 field 
trips to the southeastern United States. However, the rarer forms, which appar- 
ently have always been rare, were never encountered in fieldwork, and possibly 
no longer exist. The database of specimens is held at UWO. The taxonomic de- 
cisions have been made by the first author J.B.P) and the second author (K.D.) 
has been responsible for creating the distribution maps. 

The ultimate resolution of variation in series Parvifoliae and its putative 
interserial hybrids invites a further generation of work in whic opu- 
lation studies are supported by chromosome counts, critical morphometric 
analysis as well as molecular studies. The revisionary treatment presented here 
thus erects the platform for such work. 


TAXONOMIC TREATMENT 
series eee (Loudon) Rehder, Man. Cult. Trees Shrubs, 2nd ed.1940. sect. 


ifoliae Loudon, Arbor. et Frutic. Brit. 2.1838. Group without rank Uniflorae Beadle, nom. 
a ple Bot. Stud. 1:117. 1902. TyPE SPECIES: C. uniflora Muenchh 


Small, + xeromorphic shrubs with fine straight thorns. Leaves deciduous; usu- 
ally nearly epetiolate, petioles glandular or eglandular; blades small, longer than 
broad, entire, and often with conspicuous black glands on the tips of the teeth; 
unlobed to shallowly lobed; margins crenate or crenate-serrate; + pubescent, 
coriaceous. Inflorescences 1-few flowered, bracteolate, the bracteoles small, ca- 
ducous, linear, gland-margined, membranous to thinly herbaceous. Flowers 10- 
18mm diam.; hypanthium externally dense-tomentose; calyx-lobes foliaceous, 
us. glandular-serrate, often deeply so, as long as to somewhat longer than the 
petals; petals + circular, white; stamens 20, anthers us. ivory to cream; styles us. 
5. Fruit up to 12 mm diam., + orbicular, us. quite hairy, yellowish to ruddy; ca- 
lyx-lobes conspicuous, patent, glandular-serrate, often deeply so; nutlets 4-5, 
dorsally grooved, sides plane. 

This is a small series confined to North America. It consists mostly of one 
somewhat variable species, C. uniflora, widespread and often common in the 
coastal plain from Texas to Long Island (N.Y.) with outliers further inland and 
in Tamaulipas, Mexico. There is also a second, much rarer species, C. brittonii, 
from North Carolina and adjacent states. In addition there are three dubiously 
placed entities, C. croomiana, C. grossiserrata and C. choriophylla that will re- 
ceive brief discussion. Of the 15 names assigned by Palmer (1925) to series 
Parvifoliae (as natural group Uniflorae), one, C. choriophylla, may not belong 
and is therefore only treated briefly here. This is a very rare and probably ex- 
tinct Florida endemic tree with distinctly petiolate, thin, strongly toothed, of- 
ten slightly lobed leaves, 3-5 flowered inflorescences, cream anthers and or- 
ange-red fruit. Sargent (1922) doubted that C. choriophylla really belongs in 
series Parvifoliae,a doubt which Iam endorsing, in which case it is a rare local 
species or hybrid. Crataegus X vailiae, a putative interserial hybrid with series 
Macracanthae, is treated here for convenience, on account of its foliaceous ca- 
lyx-lobes. 

Small stature, rather glandular plants, small + unlobed leaves, often 
uniflorous inflorescences, rather small flowers, + foliaceous calyx-lobes as long 
as or longer than the petals and normally non-anthocyanic anthers, are fea- 
tures that characterize this series. Anther color may in practice be described as 
white, ivory or cream, representing both degree of precision in observation and 
some natural variation. 

Series ee is pomeeS me closely related to series Triflorae, and at 
least in istic that flowering shoots are leafy 
shoots of the season and not permanent woody short shoots. However, the plant 


426 BRIT.ORG/SIDA 22(1) 


parts are so much smaller than those of C. triflora, that even casual confusion 
is not possible. It is actually with smaller members of series Lacrimatae that 
confusion is most frequent. Members of both series are small to medium-sized 
xeromorphs that are rather glandular, often hairy and often have yellowish fruit. 
Series Lacrimatae, however, may be immediately distinguished by their zigzag 
twigs, non-foliaceous calyx-lobes much shorter than petals and fewer styles and 
nutlets. 


KEY TO SPECIES 


1. Plants often only 1-2 m tall; leaf-blades + elliptic, 1.6:1 or narrower, unlobed; an- 

thers white to crea 1.C. uniflora 
Plants 3—4 m tall at ears leaf-blades broad-ovate or broad-obovate 
“broader larger leaves usually distinctly, though sometimes only are aoe 
anthers anthocyanic 
2. Lobes of leaves obtuse; margins crenate; sides of nutlets smooth 2.C. brittonii 
2. Lobes of leaves sharper; margins with sharp, distinct teeth; sides of nutlets + 
roded 


3.C. Xvailiae 


1. Crataegus uniflora Muenchh., Der Hausvater 5:147.1770; non C. ae ord Du 
Roi, Der Harbkesche Baumzucht, ed. 1184. 1771. (Figs. 1, 2, 3, 4, 7). Tyre 
U.S.A. GEORGIA. Crawford Co.: ca. 2 mi SW of Reber: 13 Apr 2006, J.B. Phipps ee YPE 
designated here: UWO; ISONEOTYPES: A, BRIT, GA, MO, NCU, US). Muenchhausen’s original 
material would be at LINN according to TL-2 but none could be located of this species. 

Crataegus parvifolia Aiton, Hort. Kew. 2:169. 1789. Type: Specimen labeled ‘C. tomentosa Linn. 
with ‘tomentosa’ crossed out; no further information except a ‘type’ label (LECTOTYPE desig- 
nated here: Banksian Herb. BM 

a parvifolia var. flor oe Loudon, Arbor. Frut. Brit. 2:42 (no Loudon herbarium exists 

and this variety has not been neotypified; see discussion). 

cee parvifolia var. grossulariaefolia Toudon, Arbor. Frut. Brit. 2:842 (this Loudon variety 

has also not been pear see discussion). 

? Crataegus bisulcata Ashe, Bull. No iene Exp. Stn. ae 1900. 

Crataegus earlei Ashe, Bull. nee Carolina Exp. Stn. 175:112. 1900. 

Crataegus grossiserrata Ashe, Bull. North Carolina Exp. Stn. ne 112. 1900. 

Crataegus raleighensis Ashe, Bull. North Carolina Exp. Sm. 175.112. 1900. 

Crataegus armentalis Beadle, Biltmore Bot. Stud. 1:117. 1902. TYPE: U.S.A. ALABAMA Marshall Co.: 
near Albertville, no date, T.G. Harbison 3281, fruiting specimen (LECTOTYPE designated here: 


US). 

Crataegusgregalis Beadle, Biltmore Bot. Stud. L:118. 1902. Type: U.S.A. NORTH CAROLINA. Buncombe 
Co. flats of ae as River, 18 Sep 1900, Biltmore Herb. 1270b, in part (LECTOTYPE desig- 
nated here: US 

crates penance ee Elisha ane Sei. Soc. 19:12. 1903. 

Crataegus smithii Sarg, Trees & Shrubs 2 903. TYPE: U.S.A. PENNSYLVANIA. Delaware Co.: Lownes 
Run, Spr ingfield, 21 May 1902, B.H. Smith 201 (LECTOTYPE designated h 

Crataegus trianthophora Sarg,, Trees & Shrubs 2:11. 1907. TyPE: U.S.A. MISSOURL Ae Co. near 
G 


randin, (lectotype to be designated by Phipps et al., in press). 
? Crataegus croomiana Sarg,, J. Arnold Arbor. 3:202. 1922. TyPE: U.S.A. FLORIDA. Leon Co.: near 
Tallahassee, 15 Apr 1920, T.G. Harbison 5710 (HOLOTYPE: A). 


Fic. 1.T eee bi | £¢ oar gore 4) k ° fap Pies | 1 i] £ bf amar £ € ith 1350 


(MO, UWO); flowering st 1 flower details from St 214 (TENN). S. Laurie-Bourque del.; scale bars = 1 cm. 


428 BRIT.ORG/SIDA 22(1) 


— 


kK + rae ad ai 
7 J Ms P J 7 a 
from Chapman s.n. (US). S. Laurie-Bourque del.; scale bars = 1 cm. 


429 


Fic. 3. Parvifolia- or trianthophora-like form of C. uniflora showing several-flowered inflorescence form with pointed 
leaves; from Murrilf 7 (MQ). S. Laurie-Bourque del.; scale bars = 1 cm. 


Shrubs, us. under 2 m tall at maturity, less commonly to 3 m and occasionally 
reaching 5 m; extending shoots densely appressed-pubescent; one-year old twigs 
brown, older grayish; thorns 3-5(-8) cm long, usually plentiful, + straight, 
needle-like, occasionally absent, black at | yr old; young thorns commencing 
as soft, rapidly growing, needle-pointed-shoots with small linear bracts, often 
curved at first and then straightening out (Fig. 2). Leaves deciduous; petioles 
very short to lacking, glandular or not; blades (1-)1.5-3(-6) cm long, narrow- 
elliptic, elliptic, obtrullate, spatulate, oblanceolate or narrowly rhomb-elliptic, 
occasionally suborbiculate; cuneate at the base, obtuse to broadly acute at the 
tip; entire; margins crenate to dentate, with variably caducous black gland- 
tipped teeth though sometimes eglandular even young; venation semi- 


430 BRIT.ORG/SIDA 22(1) 


Ss SS \\ JB VES 


\\ 
Witen 
ei Ave 


Fic. 4 VU 4 | L tyy POF a +t 9g be 9] lat L Ef g ‘3 hd = L J] f L p ll lnb A 

leaves with veins to si tension shoots. From left hat broad-leaved form—Phipps 7677 (UWO); pos- 

sible arenicola form—Shinners 12651 (GA); C. croomiana form—Murrill in FLAS 34624; C. grossiserrata form with two 
tension st | prigt Bozeman 7188 (NCU) §. Laurie-Bourque de I.; scale bar 


431 


camptodromous, lateral veins 3-4(-5) per side, + impressed; densely scabrous- 
pubescent above especially when young, below + densely pubescent on the 
veins, thinly to moderately pubescent on the surface; often somewhat glossy 
above at maturity, + coriaceous. Inflorescences 1-3(-5) flowered; pedicels pu- 
bescent, bearing us. one or two small, caducous, linear, membranous to herba- 
ceous, gland-margined bracteoles. Flowers 10-15 mm diam:; penta 


] ns a 


strongl externally, sometimes bearing a 
linear, caducous, gland-margined bracteole; calyx- -lobes us. longer than the pet- 
als, 2-4 mm wide, + foliaceous, narrowly triangular, + deeply glandular-ser- 
rate except in rare aberrant forms, thinly pubescent on both sides; petals * cir- 
cular, white; stamens 20, anthers white to cream, except in rare, possible 
interserial hybids; styles 5. Fruit 8-10(-12) mm tall, + orbicular, tomentose, yel- 
low-green, yellow, or yellow orange to ruddy; flesh firm, dry or mealy; calyx- 
lobes to ca. 7 mm long, persistent, conspicuous, patent; nutlets 4-5, dorsally 
furrowed, laterally smooth. 

Common name.—One-flowered hawthorn. 

Distribution. —This species is widespread in the southeastern part of the 
United States (Fig. 7) from Long Island (New York) to eastern Texas with an 
outlier in Tamaulipas (Mexico). Inland it reaches the line Oklahoma, Missouri, 
southern Ohio, West Virginia. There is a remarkable disjunction in the Missis- 
sippi embayment between trans-Mississippian and cis-Mississipian populations 
that mirrors a similar pattern in C. spathulata (Phipps 1998). Crataegus uniflora 
is usually found in sand barrens and open scrubby woodland. 

Crataegus uniflora should be easy to identify on account of its small size, 
unusually large sepals (relative to the petals), uniflorous or few-flowered inflo- 
rescences and yellowish to ruddy fruit. However, it has sometimes been con- 
fused with some members of series Lacrimatae which include similar sized 
xeromorphs, also of sand-plains of the southeastern United States as is ampli- 
fied under the series discussion. Sterile plants can most easily be distinguished 
from small, sterile plants of C. berberifolia Torrey & A. Gray, a superficially 
similar member of series Crus-galli, by the smaller number of lateral veins on 
the leaves. An interesting feature of this species is the occasional deep lobing of 
extension-shoot leaves, even narrow ones, with veins to sinuses. The United 
States National Champion of C. uniflora in 1998, from Gainesville, Florida, 
measured 18 ft tall in 1992, an unusually great stature for this species. 

Crataegus uniflora is somewhat variable as befits a common species, in 
plant size, precise leaf shape (when strikingly obtrullate with acute tip or when 
suborbiculate presenting a quite different facies), number of flowers to an in- 
florescence (usually one), and anther color (usually ivory to cream). However, 
15 specific names listed under series Parvifoliae (Palmer 1925), 11 of which are 
accounted for in this paper, attest to this variation and in some cases may repre- 
sent valid, perhay tic ies or, as we shall see, perhaps interserial 


L L[ 


a°e 


432 BRIT.ORG/SIDA 22(1 


= 


hybrids. Crataegus uniflora had never hitherto been typified perhaps partly 
because the protologue is unusually diagnostic for one of the period on 
Crataegus. The protologue clearly represents C. uniflora as construed here. I have 
thus neotypified on what I regard as the widespread and common form:a small 
shrub, with coriaceous, + spatulate to narrow obovate leaves, usually numer- 
ous fine straight thorns, 1-2 flowered inflorescences, glandular-pectinate se- 
pals, anthers white to cream and yellow-green to yellow-orange fruit. Among 
this group are some particularly dwarf forms of C. uniflora from laterites in 
Georgia that have been observed with over one hundred flowers when less than 
0.3 m tall. 

There are two main kinds of deviation from the above norm, firstly, a group 
where multi (3-6)-flowered inflorescences occur and second, odd forms very 
like the putatively typical group but with often small and unlobed sepals. We 
will deal with the multiflowered types first. The most striking of those that 
can confidently be included in C. uniflora is C. parvifolia Aiton, lectotypified 
above, its type with 2-6 flowered inflorescences. A similar form is C. 
trianthophora, nicely illustrated by Faxon in Sargent’s protologue, scattered 
through the range of the species and being the main form of C. uniflora in Mis- 
souri. Multiflowered forms such as parvifolia and trianthophora represent one 
extreme of variation in the uniflora complex but the existence of two- and three- 
flowered inflorescences on some plants makes any varietal recogntion dubi- 
ous at this point. Generally, forms with multiflowered inflorescences have larger 
and relatively broader leaves than C. uniflora but this is not the case with the 
parvifolia complex. Multiflowered specimens with broad and slightly lobed 
leaves and unpitted nutlets are however, assigned to C. brittonii. 

The other main kind of variation is exemplified by C. croomiana (Figs. 4, 
9), whose type comes from Tallahassee, Florida, and which has the distinction 
of pink anthers and entire to merely glandular-serrate calyx-lobes. Out of the 
approximately 1200 specimens of the uniflora complex that in the UWO data- 
base, ten have + entire calyx-lobes and come from the northern Florida coun- 
ties of Alachua, Gadsden, Leon and Liberty. The foliage and non-geniculate 
twigs are, nevertheless, quite typical for C. uniflora. These specimens may be 
part of the variation that Murrill (1942) noticed. Further investigation is re- 
quired to see whether C. croomiana, plus perhaps some other names, should be 
upheld, possibly as sporadic interserial hybrids with members of series 
Lacrimatae. Crataegus bisulcata from North Carolina has similar calices though 
‘rounder’ leaves while the striking C. grossiserrata (Fig. 4) with a type from 
‘Florida’ (no location) also has similar calyx-lobes, though often orbicular (un- 
usual in this series) extension shoot leaves and particularly strong and sharp 
leaf-teeth. A feature of these broad, extension-shoot leaves is a tendency to deep 
lobing, sometimes with veins to sinuses. If populations of such forms could be 


433 


located today, studied and shown to be worthy of name recognition, they might 
prove to be the same as the untypified var. grossulariifolia of Loudon. 

Names listed in the synonymy but so far undiscussed all belong to entities 
more like typical C. uniflora as defined earlier and we present below a table 
illustrating some of the differences among these and other forms (Table 1). Such 
is the homonymous C. uniflora Du Roi, judging by the protologue obviously 
the same species as the Muenchhausen one, and is not worth neotypifying. The 
differences among the remainder are small and apparently not sharp, but it is 
possible that they represent local, perhaps apomictic, races. The information in 
this table is taken from protologues, amended by personal observation. Uncom- 
mon forms with suborbiculate leaves may be the same as the untypified var. 
florida of Loudon but, as these grade into typical forms, are not considered 
worthy of recognition. 


2. Crataegus pet Egel. Bull. Torrey Bot. Club 36:640. 1909. (Figs. 5, 8). Tyrr: 
U.S.A. NORTH CAROLINA. Buncombe Co.: Biltmore, 15-18 Sep 1908, W. W. Eggleston 4134 (HOLO- 
TYPE: NY; ISOTYPE: i 
? Crataegus arenicola Ashe, J. Elisha Mitchell Sci. Soc. 17:5. 1900 
? Crataegus rhodella Ashe, J. Elisha Mitchell Sci. Soc. 18:19. 1902. 


Shrub, slender, to 3 or 4 m tall; thorns 3-5 cm long, often numerous, + slender, 
+ straight to slightly recurved, blackish; extending shoots pubescent, at one 
year glabrous, somewhat shiny gray-brown, older becoming gray. Leaves decidu- 
ous; petioles 3-6 mm long, flared into the blade, pubescent; blades 2.0-3.5 cm 
long, elliptic-rhombic to broad elliptic-rhombic in general shape, generally 
widest near the middle, apex rounded to subacute, bases broad-cuneate; lobes 
O-1(-2) per side, if present, shallow, rounded: margins crenate-serrate, the teeth 
with caducous gland-tips; venation + craspedodromous, 3-4 main lateral veins 
per side; somewhat coriaceous, appressed-pubescent and dark green adaxially, 
soft pubescent and paler green abaxially; indumentum variably persistent with 
age. Inflorescences 2-6 flowered; branches dense-pubescent, bearing narrow, 
gland-bordered bracteoles. Flowers 14-18 mm diam.; hypanthium dense-pu- 
bescent; calyx lobes adaxially pubescent, narrow-triangular, margins + deeply 
glandular-serrate; stamens 20, anthers ivory; styles 3-5. Fruit subglobose, ca. 
14 mm high, red or ruddy, sometimes ripening to dull brown, pubescent; calyx- 
lobes reflexed; nutlets 3-5, lateral faces plane, dorsally ridged. 
ommon name.—Britton hawthorn. 

Distribution. —Crataegus brittoniiis known from North Carolina, Georgia, 
Tennessee and Alabama. It is found in brushy places. 

Ashe’s protologue of C. rhodella, for which no specimens were actually cited, 
states that this species comes from western North Carolina, along the Cullasagee 
River, near Franklin, Macon Co. and the Little Tennessee River, where, although 
said to be common, it has not been found since that time. Although we have 


Tasie 1. Diagnostic features of named forms in Crataegus series Parvifoliae, from protologues. 


bePy 


Stature leaf shape infl., fl. no. styleno. —fruitcolour — other 
C.arenicola shrub ovate-rhomb. 1-5 ? ruddy 
C.armentalis 0.3-1m cuneate-spatul. 1-2(-3) 3-5 d 
C. bisulcata ? rounde ? ? ry. 
C. brittonii 3-4m ovate to br. ovate 1-6 5 brown 
C.choriophylla — arborescent obov.-elliptic 3-5 4-5 or-red thornless 
croomiana 1.5- bov.-rhomb. 1 5 ? 
C. earlei shrub oblanc., obl.-cun. 1 ? ? / 
C. gregali 1-3m cun., obov.-obl. 1-2(-3) 3-5 ruddy pyriform fruit 
C. grossiserrata small obov.-spatul. ] - ? coarse leaf teeth 
C.raleighensis shrub obov.-orbic. 1-(2-3) ? yell-gr. 
C. rhodell 3m rhomb., ovate 3-6 3-5 uddy diam. fls. 18mm 
C. smithii 1-2m V 1 5-6 or.to or-gr. flowers 7-10 days 
before uniflora 

C.trianthophora 0.6-1.2m obl-obov. 3 3-4 ruddy 
abbreviations use 

br.= broad gr.= green pk. = pink 

cr= cream lacin.= laciniate ur. = purple 

cun.= cuneate obl. = oblong Serr. = serrate 

ent.= entire oblanc. = oblanceolate Spat. = spatulate 

fls. = flowers obov.= obovate wh. = white 

gl. = glandular or.= orange y.= yellow 


(L)ZZ VIS/DYO'LINE 


r n 


Fic. 5 


Biltmore Herb. s.n. ( 


si eahil ans, Oe eR 
+ sie 
DOV). S. Laurie-Bourque del.; scale 


bars = 1cm. 


ting from Small s.n. 


= 


NY); flowering from 


436 BRIT.ORG/SIDA 22(1) 


located none of this material, the protologue is sufficiently detailed that C. 
rhodella is likely conspecific with C. brittonti, the only significant difference 
appearing to be anther color (rose in C. rhodella) and, if conspecific, C. rhodella 
would become the correct name for the species. Because neither the type of C. 
rhodella, nor any authentic material annotated by Ashe, can be located, however, 
neotypifying this species is problematical. Two flowering specimens collected 
by Harbison (Harbison 11589 and 14999) from Dillsboro, Jackson Co., North 
Carolina are labelled Crataegus rhodella and with such a prominent collector 
there is necessarily some presumption that he was correct and that their anthers 
were pink. However, the label data is silent on anther color and the leaf form 
resembles C. x vailiae. Pink-anthered brittonii-like forms should nevertheless 
be carefully sought out by those that have the resources to do so. Crategus arenicola 
appears to be a smaller form of C. rhodella but in the absence of authentic ma- 
terial and similar difficulties surrounding neotypification one cannot be sure. 
A final group of larger plants (also 3-4 m tall) with longer petioles, ovate to 
broad-ovate leaves with usually sharply toothed margins, also deeply glandu- 
lar-serrate sepals and somewhat eroded nutlets is sufficiently different that they 
are referred to the putative interserial hybrid, C. x vailiae Britton, below. 

3. Crataegus Xvailiae Britton, Bull. Torrey Bot. Club 24:53. 1897. (Figs. 6, a 


TYPE: U.S.A. VIRGINIA: banks of Roanoke River, 29 May 1890, A. Brown, s.n., T. Hoge, A.M. \ 
M. Timmerman, N.L. Britton and E.G. Britton (LECTOTYPE designated here: NY). 


Crataegus missouriensis Ashe, Bull. North Carolina Exp. Stn. 175:100. 1900. TyPE: U.S.A. MISSOURI. 
Shanon Co.: Montier (NEOTYPE to be designated by Phipps et al. in press 

Crataegus conjungens Sarg., J. Arnold Arbor. 3:200. 1922. Typr: U.S.A. VIRGINI A. Alleghany Co. 
Clifton Forge, 3 May 1919, T.G. Harbison15078 (-9) (HOLOTYPE: A; ISOTYPE: NCU). 


Bushes to 2-4(-5) m tall; thorns to 2.5 cm long; extending twigs densely pubes- 
cent, at | yr thinly pubescent; at 2 yrs glabrous, dark, shiny blackish-brown to 
dark gray; older dark gray; mature bark not recorded; thorns absent or 4-6 cm 
long, few, straight to slightly recurved, somewhat narrow, black at two years. 
Leaves deciduous; petioles 3-6 mm long, pubescent, occasionally glandular; 
blades 4-5(-6) cm long, elliptic to rhomb-elliptic in general shape, symmetri- 
cal about the middle but sometimes either end widest; subacute at the tip, ta- 
pered toa cuneate base; with up to 3 extremely shallow lobes per side; margins 
with large acute or subacute teeth except near the cuneate base; margins 
eglandular (on material seen by me), teeth normally sharp; venation craspedo- 
dromous with 4-6 veins per side; appressed pubescent above, more obscurely 
so at maturity, very thin pubescent (glabrescent) on the surface below but veins 
below persistently dense pubescent; somewhat coriaceous at maturity. Inflo- 
rescences 2-8(-12) flowered; branches very dense appressed-pubescent, bear- 
ing a few caducous, linear, membranous, gland-margined bracteoles or larger 
(to 15 x 3mm) more herbaceous bracteoles in the lower parts; late-flowering. 


Fic. 6 Crataegus 
fees er eee 


ply 
1885 (A); fl ing from Harbison 15071 (NCU). S. Laurie-Bourque del.; scale bars= 


1cm. 


438 BRIT.ORG/SIDA 22(1) 


rie —— 
fe ae 4 
7 a 
fy 
SY ) 
Oe 
ae 

Number of Records 

i 0 
Fic.7 | distrit f C. uniflora in tt I United $ based on 1160 records. Not mapped 


north of Ohio River or Maryland and Delaware. 


Flowers 15-20 mm diam.; hypanthium densely spreading pubescent; calyx- 
lobes 6-8 mm long, narrowly triangular, somewhat foliaceous, abaxially slightly 
pubescent, margins deeply glandular-serrate; petals + circular, white; stamens 
20, anthers cream, pink or red; styles 3-5. Fruit 7-9 mm thick, subglobose, hairy, 
straw yellow to ruddy at maturity in cultivation at Kew, recorded as deep red 
and succulent in Missouri (Bush); calyx-lobes long, margins + pectinate, re- 
flexed in fruit; nutlets 3-5, dorsally grooved, laterally usually shallowly pitted. 
Common name.—Vail’s hawthorn. 


TERS POE 
par ne 
ApS igen | Sa 


Se Oy 


 Mumiber of Records 


iti a 3 
[ * | Literature Record 
Fic. 8.C level distributi p of C. brittonii based on 17 records seen and including literature references for C. 


sedelle faa Macon Co., NC. 


Distribution.—Vail’s hawthorn has a sparse distribution in the United States 
and is primarily a species of southern Appalachia in Virginia, Tennessee, North 
Carolina and Georgia. It is also known from Missouri and Arkansas. It occurs in 
brushy places or on riverbanks. 

With its similarities to C. uniflora, C. x vailiae may be a hybrid between 
that species and an no in series Macracanthae, most likely C. calpodendron, 
on account of its laterally eroded nutlets and late anthesis. Crataegus conjungens 


appears to be a smaller form of C. x vailiae. 

Vail’s hawthorn is an attractive plant in flower and fruit when well grown, 
especially in forms with reddish fruit, the color nicely set off by the glossy foli- 
age. It is named for Anna Murray Vail, librarian at the New York Botanic Gar- 
den, who was one of the collectors of the type material. 


440 BRIT.ORG/SIDA 22(1) 


Number of Records 
0 


Fic. 9. County level distributi fC jana based on 16 records. 


APPENDIX: FURTHER CITED SPECIMENS 


Crataegus Xvailiae Britton 
ARKANSAS. Hempstead Co.: 1905/04/21, Bush, B.F 14 (HUH).GEORGIA. Baldwin Co.: 1914/05/06. 
Harbison, 1.G. ey (HUH). Morgan Co.: 1978/05/06, Hill J.W.406 (FLAS). MISSOURI. R Aliebes 1892/ 
07/23, MacKenzie, K.K. 400 (NYBG). Shannon Co.: Hee 7, Bush, B.F. 3F (CM). NORTH CAROLINA 
Burke Co.: 1957/06/11, Bell, C.R.8759 (NC). Guilford Co.: 1927/06/29, Manning, WE.and a Ga 
1383 (HUH). Jackson Co.: 1901/05/23, Ashe WW. 32 ae Lincoln Co.: 1957/06/11, Bell, CR. 8891 
(GA).Macon Co.: 1919/05/10, Harbison, TG. (A). Richmond Co.: 1954/05/12, Melvin, L. (NCU).Rowan 
Co.: 1904/05/17, Beadle C.D. 10103 (NYBG). Swain Co.: a eee ne i I.G.1591 (A). Vance Co.: 
1874/04, Faxon, C.E. (MO). Wake Co.: 1938/04/25, ee : ‘: 3758 (MQ). VIRGINIA. Alleghany Co.: 
1909/05/19, Eggleston, W.W. 4378 (NYBG). Amelia Co.: /07/24, me vee (VCU). Bucha- 
nan Co.: 1909/10/21, Eggleston, WW. 5594 (NYBG), Sti Co.: oo Brown,H.A.et al (NYBG). 

WEST VIRGINIA. Summers Co.: 1933/05/13, Boone W. 564 (WVU). 


ee 


441 


ry 


Number of Records 
0 


=a OF CO 


1- 
ro 
9- 


4 


Fic. 10. County level distribution map of C. X vailiae based on 39 records. 


Crataegus brittonii Egg). 

GEORGIA. Rabun Co.: 1893/08/04, Small, .K. (NY). NORTH CAROLINA. Buncombe Co.: 1934/05/ 
26, Palmer, E.J.42550 (NY).Madison Co.: 1908/09/21 -22, Eggleston, W.W.4162 (NY).Stanley Co.: 1909/ 
10/24, Eggleston, W.W. 5615 (NY). TENNESSEE. Cocke Co.: 1897/08/24, Kearney, -H., Jr.596 (NY). 


Crataegus uniflora Muenchh. 

ALABAMA. Bibb Co.: 1977/04/14, Sessler, A.H. 1004 (ALU). Blount Co.: 1966/06/28, Clark, R.C. and 
Hare, T.B. 4077 (NCU). Bullock Co.: 1967/07/19, Clark, R.C. 16433 (NCU). Chambers Co.: 1967/06/02, 
Clark, R.C. 13470 (NCU). Choctaw Co.: 1964/04/20, McDaniel, S. 4222 (IBE). Coffee Co.: 1967/07/04, 
Clark,R.C. 15618 (NCU). Covington Co.: 1967/06/20, Clark, R.C. 14402 (NCU).Crenshaw Co.: 1931/04/ 
16, Palmer, J.P. 38686 (MO). Cullman Co.: 1891/06/13, Sudworth, G.B. (US). Dallas Co.: 1920/04/19, 
Harbison, T.G. 15275 (NCU). DeKalb Co.: 1955/05/06, Churchill, LA. (MSC). Escambia Co.: 1969/04/18, 
Harmon, C. 046 (NCU). Greene Co.: 1966/08/06, Clark, R.C. 6954 (NCU). Hale Co.: 1966/06/07, Will- 


442 BRIT.ORG/SIDA 22(1) 


iams,N.337(ALU).Henry Co.: 1966/07/28, Clark, R.C.6141 (NCU). Jackson Co.: 1899/06/18, Harbison, 
G. (NCU). Jefferson Co.: 1899/06/02, oe ).Lee Co.: 1897/08/11, Farle, FS. (MO). ewe 
Co.: 1982/10/03, Gunn. S.C. 1366 (AUA).M n Co.: 1969 ss Freeman, - 695 (AUA). Marne 
Co.: 1967/06/05, Clark, R.C. 13644 (NCU). Sie Co.: 1977/04/23, Ross, L. 107 (ALU). Mobile Co. 
1898/04/15, Mohr, C. (ALU). Monroe Co.: 1959/06/1 es en,H. ne rnenee Co.: 188 0/07/ 
15, Mohr, C. (US). Perry Co.: 1967/08/10, Clark, R.C. 17835 (NCU). Pike Co.: 1931/04/15, Palmer, EJ. 
38656 (MO).Randolph Co.: 1967/04/16, Clark, 2.C. 11365 (NCU). St. Clair Co.: 1951/06/24, Hollis, C.G. 
(ALU). Sumpter Co.: 1966/06/06, Drapalik, D. 1362 (NCU). Talladega Co.: 1966/06/20, C oe R. 5 
Heard, 1A, 2895 (NCU). Tallapoosa Co.: 1973/10/06, Pringle G. ie aa A). Tuscaloosa Co.: 1982/1 
07, Burckhalter, R.015 (UNA).Walker Co.: 1967/05/08, nee A.C. 12564 (NCU).Winston Co.: ie /05 
10, Clark, R.C. 12995 (NCU), ARKANSAS. Cleburne Co.: 1967/0 nos Demaree, D. 56271 (IBE). Dallas 
Co.: 1975/07/05, Sparkman 69902 (IBE). ast Co.: a 0/13, Palmer, EJ. 26535 (A). Fulton Co 
1909/05/21, Bush, B.F. 5690 (A). Garland Co.: 1925/04/23, Palmer, EJ. 26885 (A). Howard Co.: 1909/ 
10/07, Kellogg, JH. (A). Independence Co.: | on Thomas, D, 9093 (NLU). lzard Co.: 1929/04/ 
27, Palmer, EJ. 35547 (A). Marion Co.: 1914/06/14, Palmer, EJ. 5974 (A), Quachita Co.: 1975/08/06, 
Demaree, D. 69628 (\BE). Pope Co.: 1932/09/20, Merrill, G. 36 (A). Saline Co.: 1972/07/15, Demaree, D. 
6524] (IBE). Sharp Co.: 1970/07/24, ali D. 15255 (NLU). Stone Co.: 1971/09/18, Demaree, D.64280 
(IBE). White Co.: 1887/07/23, Coville FV. 171 (US). DELAWARE. Kent Co.: 1941/05/09, Tatnall, R.R. 
» (DOV). New Castle Co.: 1930/05/30, - nall, R.R. 896 (DOV). Sussex Co.: 1902/09/12, Norton, 
J.B.S. (MARY). District of Columbia: 1889/05/18, Burgess, E.S. (MARY).FLORIDA. Alachua Co.: 1982/ 
03/14, Easterday, J.C. 821 (USF). Clay Co.: 1869/03, Canby, WM. (US). Columbia Co.: 1895/07/11-19, 
Nash, V. (A). Duval Co.: 1894/07/09, Curtiss, A.H.4553 (US), Escambia Co.: 1949/04/15, Hood, S.C. 1915 
(FLAS). Franklin Co.: 1923/09/27, Harbison, 1.G.6046 (A).Gads 1966/04/30, McDaniel, S. 7407 
(IBE). Hamilton Co.: 1900/04/17, Hugar, A. pl Jackson Co.: 1929/04/12, Palmer, E.J. 35299 (US). 
Jefferson Co.: 1950/09/30, Kruz, H. (FSU). Leon ioe 1920/10/06, re nison, 1.G.5710 (A). Liberty Co.: 
1957/05/10, Uttal, JL. 5382 (LYN). Marion a 85/07/18, Hansen, B.F, Van Etten, M. and Wunderlin 
R.P. 9981 (USF). Polk Co.: 1985/06/12, Hopkins, : 3410 (FLAS). Union Co.: 1939/08/18, Murrill, WA. 
(FLAS).Wakulla Co.: 1964/06/21, Godfrey, R.K.64178 (FSU).GEORGIA. Bartow Co.: 1971/08/01, Wa- 
ters, J.C. 136 (AUA). Berrien Co.: 1973/05/06, McCarty, I. (MO). Bibb Co.: 1901/05/26, No Collector 
(MO). Brooks Co.: 1966/07/13, Faircloth, W.R. 3465 (MO). Burke Co.: 1938/04/10, McVaugh, R. and 
Byron, J.H. 2470 (GA). Chatham Co.: 195 ie Duncan, WH. 21113 (DHL). Clarke Co.: 1929/03/12, 
Dunnaway, JM, E7433 (GA). ee Cox 07/26, Bozeman, J.R. 6614 (NCU). Cobb Co.: 1943/05/02, 
Wurdack, J. Saale Co.: 1900/0 es Sargent, C.S. (A). DeKalb Co.: 1966/( eae, Wood, A. (GA). 
oughert « 1900/08/13 pane W.R.and baie .505 (US). Echols Co.: 1965/07/16, Dean, R. 
and Faircloth, te 2297 (GA). Elbert Co.: 1977/09/10, Coile, N.C. 1424 (GA). Flo on Co.: 1956/08/09, 
Lipps, L. 957 (TENN). Glascock Co.: 1966/07/15, Faircloth, WR. 3499 (MO). Greene Co.: need 8/13, 
Duncan, WH. 1585 (GA). Gwinnett Co.: 1908/04/12, Allard, H.A. 106 (US). Habersham Co.: 1914/05/ 
06, Ashe, W.W. (NCU). Hancock Co.: 1966/08/09, Bozeman, LR. 7314 (NCU). Hart Co.: 1942/04/26, 
Duncan, W. 4846 (GAM). Houston Co.: 1979/06/01, Dixon, D. ee (NLU). Jackson Co.: 1948/04/30, 
Steen A. 5085 (US). es Co.: 1966/08/08, Bozeman, ee 8 (NCU). Lanier Co.: 1965/07/30, 
Faircloth, WR. and Norsworthy, J. 2603 (NCU). Liberty Co.: a Harbison, I.G. (A). Lowndes 
a 1973/05/17,McCarty, I. (MO).Macon Co.: 1898/1 0/02 aie WM., Muir, J.and Sargent, C.S. (DOV). 
Meriwether Co.: 1945/04/17, Henry, M.G. 3982 (GA). Morgan Co.: 1979/04/22, Hill, L. W848 (GA). 
sgh Co.: LA ae Montgomery, F. 1025 (GA). pet Co.: 1966/07/02, Clark, LV. (GA). Put- 
«1942/04/30, Duncan, WH. 4913 (GA).Rabun Co.: 1893/08/07, Small, .K. (MO). Richmond 
oe 1900/04/25, psa 1 (DOV ). Screven Co.: Peer 7,Cron nee 5004 (GA). Stephens 
Co.: 1975/07/21, Boufford, D.E. and Wood, E.W. 2096 Meseaaans Co.: 1940/05/11, Duncan, WH. 
2342 se Troup Co.: 1829/06/12, Sargent, C.S. (A). Turner Co.: 1947/1 7 Thorne, R.F. 7509 (GA). 
Washington Co.: 1970/06/26, Hobbs, G.B. and ons " 52 (NCU). ILLINOIS. Wabash Co.: 1875, 
Schneck, i (MQ). KENTUCKY. Boyd Co.: 1923/05/20, Horsey, R.E. 1662 (A). Pulaski Co.: 1939/09/08, 


pate 


tN 
Co 
wo 
[o> 


— 


=~ 


Brown, E.L.2637 (A). LOUISIANA. Bi ille Parish: 1967/06/07, Thieret, /. W. 26568 (LAF). Caddo Par- 
ish: 1969/06/15, Thieret, J.W. 31277 (LAF). Caldwell Parish: 1973/06/02, Marx, Pet al 1207 (NLU). Clai- 
borne Parish: 1975/04/26, Haynes, R.R.4807 (ALU). Desoto Parish: 1979/11/10, Dixon, D. 2526 (NLU). 
Morehouse Parish: 1972/05/04, Thomas, D.R. 28684 (NLU).Orleans Parish: 1916/04, Cocks, R.S. (NO), 
Rapides Parish: 1976/06/18, Shultz, CJ). and S.£. 142 (NLU). Tangipohoa Parish: 1901/09/01, Cocks, 
R.S. (NO). Union Parish: 1983/07/23, Moore, D.C. 1670 (NLU). MARYLAND. Allegany Co.: 1940/06/ 
12, Norton, J.B.S. (MARY). Caroline Co.: 1928/06/03, Tatnall, R.R. 78 (DOV). Montgomery Co.: 1918/ 
09/29, Hunnewell, FW. 5661 (GH). Washington Co.: 1906/07/11, Jones, WR. and Shreve, F. 689 (US). 
Wicomico Co.: 1903/06/06, Norton, J.B.5. (MARY).Worcester Co.: 1940/06/24, Jeffery, W.F.and Norton, 
J.B.S. (MARY).MISSISSIPPI. Clarke Co.: 1967/09/09, Jones, S.B.and Jones, C. 15241 (NCU). pee Co.: 
1961/08/20, McDaniel, S. 2689 (IBE). Jones 1978/06/03, Morgan, D.678 (IBE). Kemper Co.: 1859/ 
05/01, Holgand, £. (MO).Lauderdale Co.: 1922/04/18, Harbison, T.G.6043 (A).Lowndes Co.: 1 ee 
02, Holmes, W.C. 1701 (NATC). Newton Co.: 1961/07/22, McDaniel, 5.2627 (IBE). Wayne Co.: 1967/04/ 
12, Bass, C, Jones, 5.B., Prescott, V. and Reynolds, J.D. 11631 (GA). MISSOURI. Carter Co.: 1893/05/16 
Eggert, H. (MO). Dent Co.: 1927/09/06, Kellogg, J. (MO). Douglas Co.: 1934/08/19, Steyermark, J. 14705 
MOQ). Ozark Co.: 1928/06/28, Palmer, E.J. 34812 (GH). Reynolds Co.: 1934/09/28, Steyermark, J. A. 
15577 (MO). Shannon Co.: 1978/05/18, Summers, B. 396 (MO). St. Louis Co.: 1879/09/16, eee 
G. (MO). Texas Co.: 1934/08/15, Steyermark, J.A. 14530 (MO). Wayne Co.: 1893/06/22, ae >} 
(CM). NEW JERSEY. Burlington Co.: 1955/08/18, Churchill, A. (MSC). Cape May Co.: 1 i 5, 
Killip, E.P 2406 (US). Cumberland Co.: 1964/07/27, Gillis, W.T. 5646 (MSC). Monmouth Co.: 1976/06/ 
24, Schwenk, J. (MO). Salem Co.: Vanco ip d,J.H. 1902 (MO).NEW YORK. Cattaraugus Co.: 
1895/07/17, Andrews, E.F (AUA). Richmond C 79/08/22, Schrenk, J. (MO). Suffolk Co.: 1879/06/ 
03,LSK (MO). NORTH CAROLINA. Alamance a ee Hammond,A.E.and Ramseur, G.$.2254 
(NCU). Alexander Co.: 1935/05/08, Verrhoof, O. (NCSC).Anson Co.: 1956/09/21, Ahles, H.E.and Leisner, 
R.S. 19469 (NCU). Bladen Co.: 1948/06/10, Fox, WB. et al 1705 (NCSU). Brunswick Co.: 1954/08/01, 
Melvin, L. (NCU). Buncombe Co.: 1908/09/15, Eggleston Denote ieee Co.: 1958/09/09, Bell, 
R.C. 15054 (NCU). Carteret Co.: 1923/06/30, Holmes, J.S. (NCU). Caswell Co.: 1958/05/22, Bell, RC 
17893 (NCU). Catawba Co.: 1891/06/25, Heller, AA and Small, J.K. (US). Chatham Co.: 1988/05/19 
Swab, £.C. 162 (USCH). Chowan Co.: 1898/07/29, Kearney, T. Jr. 1903 (US). Cleveland Co.: 1956/06/22, 
Ahles, H.E. and Leisner, R.S. 15253 (NCU). Craven Co.: 1919/05/26, Harbison, T.G. 15090 (NCU). 
Cumberland Co.: 1957/05/04, Ahles, H.E.and Hammond, A.24389 (NCU). Davidson Co.: 1916/06/07, 
Totten, H.R. (NCU). Davie Co.: 1916/06/08, Totten, H.R. (NCU). Durham Co.: 1932/08/02, Blomquist, H.L. 
83 (US). Forsyth Co.: 1948/09/07, Correll, D.S. and Correll, H.B. 14478 (DUKE). Franklin Co.: 1956/07/ 
10, Ahles, H.E. and Leisner, B.S. 16508 (NCU), Gaston Co.: 1956/09/17, Ahles, H.E. and Leisner, R.S. 18810 
(NCU). Granville Co.: 1956/05/22, Ahles, H.E.and Bell, C.R. 12756 (NCU). Greene Co.: 1934/05/12, Tot- 
ten, H.R. (NCU). Guildford Co.: 1953/05/12, Melvin, L. (NCU). Halifax Co.: 1956/07/19, Ahles, H.E. and 
Leisner,R.S. 16975 (NCU). Harnett Co.: 1956/09/29, Laing, H.349 (NCU). Henderson Co.: 1962/08/21, 
Pittillo, D.546 (GA). Hertford Co.: 1958/07/08, Ahles, H.E.and Duke, J.A.46018 (NCU). Iredell Co.: 1958/ 
05/18, Ahles, H.E.and Britt, R.41050 (NCU). Jackson Co.: 1940/07/1 tere H.R. (NCU). Johnston Co.: 
1919/04/26, Harbison, I.G. 14967 (NCU). Jones Co.: 1954/08/08, Corbin and pee (NCU). Lee Co.: 
1955/06/27, Kessler, P-A. 350 (NCU). Lenoir Co.: 1908/10/12 see WW. 4279 (US). Lincoln Co.: 
1957/06/11, Bell, R.C.8907 (NCU). Madison Co.: 1908/09/21, Eggleston ie: (CM).Mecklenburg 
Co.: 1958/10/02, Ahles, H.E.and Duke, .A.49968 (NCU).Montgomery Co.: 1970/05/18, Wells, E.F 2888 
(NCU). Moore Co.: 1900/08/20, Ashe W.W. (CM). New Hanover Co.: 1963/06/30, Ahles, H.E. and Mc- 
Crary, A. 537 (NCU). Northampton Co.: 1958/07/07, Ahles, H.E. and Duke, JA. 45726 (NCU). Orange 
Co.: 1969/08/08, Whigham, D. 1251 (NCU). en Co.: 1951/07/31, Batson, WT. Jr. 749 (DUKE). Pitt 
Co.: 1956/05/09, Eason, M. 20 (NCU : ae Co.: 1921/08/11, Peattie, D.C. 1163 (NCU). Randolph Co.: 
1965/10/10, Harper, FH.and Harpe een Rockingham Co.: 1956/07/07, abi F.13628 
(NCU). Rowan Co.: 1908/09/1 2 oe eston, WW. 4105 (CM). Rutherford Co.: 1956/08/30, Melvin, L. 
5007 (NCU). Scotland Co.: 1937/05/03, Coker, W.C. (NCU). Stanly Co.: 1891/08/18, Heller, A.A. and 


— 


444 BRIT.ORG/SIDA 22(1) 


Small, .K.49 (MO).Stokes Co.: 1958/07/26, Radford, A.E. 37727 (NCU). Transylvania Co.: 1961/07/05, 
Rodgers, C.L.61328 (DUKE). Vance Co.: 1956/10/01, Ahles, H.E. and Leisner, R.S.20430 (NCU). Wake Co.: 
1964/07/14, Sawyer, G.P Jr. 1256 (USCH). Warren Co.: 1968/06/12, O'Briant, S. and Radford, A.£. 45482 
(ALU).Washington Co.: 1958/06/15, Radford, A.F. 35225 (NCU). Wayne Co.: 1957/05/03, Radford, A.E. 
22009 (NCU). Wilkes Co.: 1938/06/13, Stewart, S.C. and Stewart, L. (NCU). OKLAHOMA. Choctaw Co.: 
1850s-1900s, Edwards, J.E. (GH). Garrett Co.: 1905/06/08, Shreve, F483 (US).Marshall Co.: 1963/06/ 
19, Ware, G. (NATC). Pusmataha Co.: 1931/05/26, Palmer, EJ. 39406 (MO). PENNSYLVANIA. Clinton 
Co.: 1895/06/10, Glatfelter, N.M. (MO), Delaware Co.: 1903/05/20, Graves, J.A. (MO).North Hampton 
Co.: 1891/08/03, Porter, T.C. (US). SOUTH CAROLINA. Abbeville Co.: 1957/06/29, Radford, AE. 25986 
(NCU). Aiken Co.: 1898/08/06, Eggert, H. (MO). Allendale Co.: 1953/04/06, Batson, W.T.and Kelley, WR 
(USCH). Anderson Co.: 1809/07/24, Davis, J. (CM). Barnwell Co.: 1952/06/13, Batson, W.T.and kelley, 
WER. (USCH). Beaufort Co.: 1917/04/29, Churchill, LR. (MO).Calhoun Co.: 1957/05/19, Ahles, H.E.and 
Haesloop, J.G. 25520 (NCU). Charleston Co.: 1957/05/20, Ahles, H.E. and Haesloop, J.G. 25769 (NCU). 
herokee Co.: 1957/09/18, Ahles, H.E. 34428 (TENN). Chester Co.: 1957/06/27, Bell, R.C. 9536 (NCU). 
Clarendon Co.: 1914/05/22, Stone, W. 271 (A). Colleton Co.: 1983/04/04, Rayner, DA. 1593 (USCH). 
Darlington Co.: 1941/05/15, Smith, B.£. 947 (USCH). Edgefield Co.: 1957/05/12, Radford, A.E. 22648 
(NCU). Fairfield Co.: 1983/06/22, Gould, B., LaBorde, O.and Pugh, A.E. (NLU). Florence Co.: 1982/04/ 
24, McFarlane, 8.1191 (USCH). Greenville Co.: 1934/09/20, Wherry, E.7. (A). Horry Co.: 1957/09/12 
Totten, H.R. 1576 (DHL). Jasper Co.: 1954/04/19, Ahles, H.F. 7807 (NCU). Kershaw Co.: 1960/05/25, 
Herring, WA. 3 (USCH). Lancaster Co.: 1975/05/10, Boufford, D.E. 15999 (CM). Laurens Co.: 1991/07/ 
09, Horn, C.N. and Mytko, D.E.4564 (USCH). Lee Co.: 1957/06/06, Radford, A.E. 24320 (NCU). Lexington 
Co.: 1934/05/19, Palmer, E.J.42423 (MO).McCormick Co.: 1960/10/14, Mc Comb, C.65 (GA).Newberry 
Co.: 1957/06/25, Bell, R.C. 9186 (NCU). Oconee Co.: 1942/05/09, Prince, A.E. 4001 (A). Orangeburg 
Co.: 1983/05/23, Aulbach-Smith, CA, Long, G.and Tallevast, T. 2476 (USCH). Pickens Co.: 1975/07/18, 
oe . and Wood, E.W.2077 (NCU).Richland Co.: 1976/08/26, Ne/son, /.B.68] (IBE). Spartanburg 
06/04, Bell, R.C.8169 (NCU). Sumpter Co.: 1933/04/17, Harbison, I.G.and Totten, H.R. (NCU). 
aioe ea 993/05/09, Horn, C.N. 6627 (USCH). Williamsburg Co.: 1957/06/12, Radford, A.£. 24830 
(NCU). York Co.: 1957/06/05, Ahles, H.E.and Haesloop, .G. 27098 (NCU). eee aera Co.: 
1961/06/30, Ellis, WH. 28599 (TENN). Cocke Co.: 1898/07, Ruth, A. 286 (US). Hamilton Co.: 1921/04, 
eet 1024 (US).Knox Co.: 1938/06/29, Sharp, AJ. 232 (TENN).Lawrence Co.: 18 ea 8, Ruth, 
4 (MO). Union Co.: 1934/07/10, Kelly 2473 (TENN). TEXAS. Bell Co.: 1902/05/08, Reverchon, J. 
a Jasper Co.: 1963/04/14, Correll, D.S. a Correll, H.B.27229 (MO).Smith Co.: 1949/05/15, Shinners, 
JH.11183 (GA). VIRGINIA. Alleghany Co.: 1909/05/19, Eggleston, W.W.4374 (MO).Amelia Co.: 1937/ 
05/25, Lewis, J.B. 555 (VPI). Amherst Co.: eas 1, Crandell, D. 9669 (LYN). Appomatox Co.: 1967/ 
07/07, James, FC. 7231 (NCU). Bedford Co.: 1871/08, Curtiss, AH. (GH). Botetourt Co.: 1967/05/06, 
James, F.C. (NCU). Brunswick Co.: 1968/04/21, James, F.C. 13252 (NCU). Buckingham Co.: 1951/06/ 
29, Jarrett, G.P and Little R.L. 14622 (US). Campbell Co.: 1893/07/20, Hel pone 099 (MO). Caroline Co.: 
1968/04/28, James, FC. 13895 (NCU). Chesterfield Co.: 1971/06/10, Johnson, M.F.23220 (NCU). Craig 
Co.: 1967/07/19, James, EC. 7630 (NCU). Dinwiddie Co.: 1965/08/18, James, FC. 2375 (NCU). Fairfax 
Co.: 1970/07/31, Kiltz, BF. 57 (NLU). Fluvanna Co.: 1921/06/03, Eggleston, WW. 17740 (US). Franklin 
Co.: 1966/07/15, Ramsey, G.W.et al 4938 (LYN). Frederick Co.: 1941/06/19, Palmer, E.J.45271 (A). sas 
Co.: 1935/09/07, Adams, .W.and Wherry E. 2323 (A). Goochland Co.: 1970/05/28, Johnson, M.F. 320 
Snes ae Co.: 1944/06/22, foe M.L. and Lewis, J.B. 14609 (US). Hanover Co.: 1 se 
26, James, F.C. 13446 (NCU). Henry Co.: 1968/05/28, James, F.C. 10443 (NCU). Lee Co.: a 
James, FC.9791 ae Louisa Co.: ieee Sharp, AJ. 1102 (TENN).Lunenburg Co.: 1965/09/19, 
James, FC. 3071 (NCU). Mathews Co.: 1852/05, Bischoff, G.G. (MO). Mecklenburg Co.: poner 
Seaman, a (NCU). Middlesex Co.: 1966/07/14, James, F.C. (NCU). Montgomery Co.: 1962/ 
07/02, Kral,R 8 (FSU).New Kent Co.: 1977/05/12, Bims, S. (VCU). Patrick Co.: 1950/07/24, van 
B.6813 ae ae Co.: 1893/07/30, Heller, A. 1099 (A). Powhatan Co.: 1967/06/08, jeniee 
F.C.6279 (NCU). Prince George Co.: 1936/06/19, Fernald, M.L.etal 5791 (GH).Roanoke Co.: 1969/07/ 


— 


445 


29, Uttal, L.J.6648 (VPI). Rockbridge Co.: 1881/05, /.8.C. (MO).Rockingham Co.: 1956/07/01, Allard 
H.A. 21957 (US). Shenandoah Co.: 1970/05/19, Churchill, A. (MSC). Smyth Co.: 1892/06/20, Small, 
JK. (US). Suffolk City, 1980/09/17, Wieboldt, TF. 590 (WILLI). Surry Co.: 1934/08/17, Totten, H.R. (NCU). 
Zuni Co.: 1908/08/19, Render, A. (A). WEST VIRGINIA. Grant Co.: 1979/05/15, Duppstadt, H.029 (WVA). 
Greenbrier Co.: 1959/07/29, Clarkson, R.B. 2992 (NCU). Hampshire Co.: 1929/05/30, Walker, EH. 303 
(GA). Hardy Co.: 1955/06/16, Bartley, F. and Hicks, L.E.2122 (NY). Mineral Co.: 1928/10/06, Saat 
W.W. & Kelly, J.W. 22859 (US).Morgan Co.: 1969/05/10, Downs, R.M. 6349 (NCU). Wyoming Co.: 1900/ 
08/13, Morris, E.L. 7189 (US). 


ACKNOWLEDGMENTS 


The curators of the following herbaria are thanked for the loan of specimens: A, 
ALU, AUA, BALT, CM, DHL, DOV, DUKE, FLAS, FSU, GA, GH, IBE, JSU, LAF 
LSU, LYN, MARY, MO, MSC, NATC, NC, NCC, NCSC, NLU, NO, NY, TENN, UCC, 
UMO, UNA, US, USCH, USF, VCU, VPI, VSC, WCU, WILLI, WVA. Vicky 
Papworth (BM) went to great lengths to help with the type of C. parvifolia. Bob 
O’Kennon of the Botanical Research Institute of Texas, Fort Worth and Ron 
Lance, now of Chimney Rock Park, North Carolina, are thanked for their par- 
ticipation in some of the first author's field trips to the area. Ms. Alison Lacon, 
who was an undergraduate at the time, is thanked for compiling the large speci- 
men database of C. uniflora without which the detail of mapping used here 
could not have been attained. Susan Laurie-Bourque is thanked for the beauti- 
ful line illustrations. The National Sciences and Engineering Research Council 
of Canada funded this work via a discovery grant to the first author. 


REFERENCES 


Cronauist, A. 1980. Vascular flora of the southeastern United States, vol.1, Asteraceae. Uni- 
versity of North Carolina Press, Chapel Hill. 

Murritt, W.A. 1942. New Florida hawthorns. Castanea 7:19-30. 

Pavmer, E.J. 1925. Synopsis of North American Crataegus. J. Arnold Arbor. 6:5—128. 

Puipps, J.B. 1988. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. In- 
troduction and series Aestivales. J. Arnold Arbor. 69:401-431. 

Puipps, J.B. 1998. Synopsis of Crataegus ser. Apiifoliae, Cordatae, Microcarpae and Brevispinae 
(Rosaceae subfam. Maloideae). Ann. Missouri Bot. Gard. 85:475-491. 

Pipes, J.B., R.J. O’KeENNoN, and R.W. Lance. 2003. Hawthorns and Medlars. Timber Press, Port- 
land, OR. 

Pripps, J.B.,G. Yarskievycn, and K.Woop.(in press). Typification of Crataegus (Rosaceae) names 
from the Missouri flora. Harvard Pap. Bot. 

Sarcent, C.S.1922.Notes on North American trees, X:new species and varieties of Crataegus. 
J. Arnold Arbor. 3:182—207. [Crataegus choriophylia, p.201] 


BRIT.ORG/SIDA 22(1) 


Book REVIEWS 


SHatolyA Dr La Tour and RichArp De LA Tour. 2001. The Herbalist’s Garden: A 
Guided Tour of 10 Exceptional Herb Gardens: The People Who Grow Them 
and the Plants That Inspire Them, (ISBN 1-58017-410-8, hbk.). Storey Pub- 
lishing, 210 MASS MoCA Way, North Adams, MA, 01247. U.S.A. (Orders: # 
67410, 1-800-441-5700; wwwstorey.com). $27.50, 229 pp. color photos, color 
illustrations, 8 3/8" x 10" 

Herbalists and gardeners (or both) will enjoy this very readable coffee ele book. The authors 

vho also run an herb store 


are a husband-and-wife team of passionate medicinal plant gardeners 
This book is a tour of the gardens of ten well-known herbal medicine experts ai tas une 


Rosemary Gladstar, and others. Each section includes - Ey shies of the BarccnErs and the h 


| 
OLLIE LO really Step inside thes Se Calill 


tory behind their gardens. The copious color 
retreats. A particular gem is the section on James | Duke’ s “farmette” in Maryland. A renowned ethno- 

botanist and researcher, Dr. Duke's six-acre herb garden includes 80 plots, each centered ona specific 
health condition, such as cancer or respiratory conditions. Included in each section are lists of the 
gardeners’ favorite plants, watercolor garden plans, medicinal tea recipes, and tips for growing your 
nnician, Botanical Research Institute of 


own medicinal plants.—Marissa Oppel, MS, Herbarium Tec 
Texas, Fort Worth, Texas 76102-4060, U.S.A. 


LONDA SCHIEBINGER. 2004. Plants and Empire: Colonial Bioprospecting in the At- 
lantic World (ISBN 0-674-01487-1, hbk.). Harvard University Press, 79 Garden 
Street, Cambridge, MA 02138, U.S.A. (Orders: www.hup.harvard.edu/ 
index.html). $39.95, 320 pp., 20 black-and-white illustrations, 6 1/2" x 
91/2" 


This page-turner is a fascinating glimpse into the mone of colonial! pa eats duri ane the’ ae 
] 


LATS 


eighteenth century. ” When botany was young, voyaging 
from the New World ina mad race to discover the most economically and medicinally useful ae 


Colonial bioprospecting was a cut throat business involving botanical spies and pirates. The author 
presents an overview of this rough-and-tumble era, focusing on the voyaging botanists and those 
armchair botanists who stayed in Europe and had their plants collected by assistants, ship captains, 
and others. A botanical artist and scientist, Maria Sibylla Merian, was one of very female voyaging 
botanists during this time period. While in the rae bean, Merian described the “peacock flower” 
and its use by slave women as an abortifascient. Schiebinger’s ¢ arefully constructed history of this 


discovery and the reasons it never reached mainstream Europeans is an interesting portrait of Euro- 
pean reproductive mores and the business of botany in the 1700s.—Marissa Oppel, MS, Herbarium 
Technician, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


SIDA 22(1): 446. 2006 


ANEW SPECIES OF SIENANTAIUM 
(MELANTHIACEAE) FROM TENNESSEE, U.S.A. 


B. ie Wofford 


Dept. of Fc Biolog 
The Uni iyeisiy of nee 
SA 


Knoxville Tennessee 37996 


ewofford@utk.edu 


ABSTRACT 


Stenanthium diffusum Wofford, sp. nov, is described as a new species from Fentress, Morgan, Pickett, 
and Scott counties on the northern Cumberland Plateau of Tennessee. It is most closely related to S. 
gramineum but differs in having (1) a more diffuse inflorescence, (2) flowers on the mid-portion of 
lateral panicle branches spaced =8 mm apart and with distinct pedicels (1.6-4 mm long), and (3) the 
uppermost non-bracteal stem leaf less than 15 cm above ground level. It also flowers later and has 
ecological requirements differing significant 


y from S gramineum Stenanthitum liffus a4) 11S an ex- 


tremely rare rockhouse endemic in need of consideration for state and federal ee species 
status. 


RESUMEN 
Se describe una nueva especie Stenanthium diffusum Wofford, sp. nov., de los condados de Fentress, 
Be Hee y peek del norte ne Gumbel and Plateau de Tennessee. Esta estrechamente 


tener (1) una eee mas difusa, (2) flores en la 
parte media lateral de las ramas eidel paniculo espaciadas 28 mm y con aoa otis (1.6-4 
mim de longitud), is (3) la ba superior del tallo no Practeal a menos ede cm sobre el nivel del suelo. 
También f] y = tante S.gramineum 
Stenanthium diffusum es un eaaone ea raro de sale que requiere el estatus 
de especie amenazada estatal y federalmente. 


TAXONOMIC/NOMENCLATURAL PERSPECTIVE 


In the traditional sense, the genus Stenanthium (Liliaceae sl.) is represented by 
one species in Asia, S. sachalinensis F Schmidt, and from two to four taxa in 
North America. Stenanthium occidentale A. Gray, western featherbell, occurs 
in California, Oregon, Washington, Idaho, Montana, British Columbia, and Al- 
berta (USDA, NRCS). In the eastern United States, depending upon taxonomic 
interpretation, Stenanthium may be composed of the typical variety, S. 
gramineum (Ker Gawler) Morong var. gramineum (eastern featherbell), and two 
additional varieties, S.gramineum var. micranthum Fernald and S.gramineum 
var. robustum (S. Watson) Fernald. Stenanthium gramineum sl. ranges from 
Florida to Texas and northward to Indiana and Pennsylvania. Local established 
waifs are known from upper Michigan (Voss 1972). 

Fernald (1946) provided a treatment of the Stenanthium gramineum com- 
plex in the eastern United States and concluded that there is a a confluent series 


SIDA 22(1): 447 — 459. 2006 


BRIT.ORG/SIDA 22 


— 
= 


) 


of variation and that...“As the best Ican now do propose the following defini- 
tion of the three varieties.” Although it is my impression that he had little faith 
in their distinctiveness, he did provide the following key for others to compare 
with their collections: 
a. Stem (dry) 4-10 mm thick at lowest exposed internode; leaves rather crowded be- 
low, rapidly cnn below paniegyl firm to coriaceous, ly opaque, the larger 
one wide, t ell prominently raised {ribs ee ce urface; 
panicle lax, the ae distant, the flowers mostly chine ee the often 
flexuous branches; perianth 3-8(-10) mm long; capsules ovoid-urceolate, 6-9 mm 
long, on spreading to reflexed pedicels; seeds 5-5.5 mm ne 
Stem 0.5-1.9 m high, 4-10 mm thick at base; perianth 5-10 mm long S.gramineum 
var. typicum (= var. gramineum) 
Stem 0.25—1 m high, 1.5-5 mm. thick at the base; perianth 3-4(-5) mm long 
var. micranthum 


a. Stem (dry) 7-15 mm thick at lowest exposed internode, up to 1.8 m high; leaves 
crowded and numerous nearly up to panicle, thin and membranaceous, translu- 
cent, the larger ones 1-3 cm wide, their ribs mostly immersed in the tissue; panicle 
usually dense, with flowers crowded along the stiffly ascending branches; perianth 
5-10 mm long;capsules oblong-subcylindric, 9-10 mm long, crowded and ascend- 
ing to horizontally spreading; seeds 5-8 mm long 

Since Fernald’s (1946) assessment, the two varieties of typical S. gramineum 

have been largely unrecognized as distinct taxa in local and regional floras of 

the eastern United States as well as in the recent Flora of North America treat- 
ment by Utech (2002). Utech also cited and agreed with Gates (1918) and 

Johnson (1969) that the varieties of S.gramineum are indistinct and sympatric. 

lagree that significant morphological variation and overlap of characters exist 

but the extremes, however, are strikingly distinct and warrant consideration 
for additional study. 

Recently, Zomlefer and Judd (2002) made the following significant new 
combinations in the tribe Melanthieae based on parsimony analysis of ITS and 
trnL-F DNA sequence data, distributions, chromosome numbers, and morpho- 
logical data: Stenanthium sachalinensis = Anticlea sachalinensis (F. Schmidt) 
Zomlefer & Judd, Stenanthium occidentale = Anticlea occidentalis (A. Gray) 
Zomlefer & Judd, Zigadenus densus (Desr.) Fernald = Stenanthium densum 
(Desr.) Zomlefer & Judd, and Zigadenus leimanthoides A. Gray = Stenanthium 
leimanthoides (A. Gray) Zomlefer & Judd. These former Zigadenus taxa share 
significant morphological and geographical distinctions from the S.gramineum 
complex and their transfer to Stenanthium may be subject to debate by bota- 
nists emphasizing morphology and biogeography. Both Zigadenus densus an 
the poorly defined and taxonomically ambiguous Z. leimanthoides have a small 
gland at the base of the obovate, round-tipped tepals. They are primarily re- 
stricted to the southeastern U.S. Coastal Plain with the exception of a few out- 
liers in the mountains of West Virginia, Virginia, and North Carolina, and on 
the Eastern Highland Rim of Tennessee. Stenanthium gramineum and S. 


var. robustum 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 449 


diffusum lack a gland at the base of the lanceolate, tortuose-tipped tepals. 
Stenanthium gramineum is widespread throughout most of the eastern U.S. 
while S. diffusum is restricted to the northern Cumberland Plateau of Tennes- 
see. Conversely, adoption of the new combinations of Zomlefer and Judd would 
leave the genus Stenanthiumss. restricted to the eastern United States. 


BIOLOGICAL/ ECOLOGICAL / FLORISTIC RELATIONSHIPS 


The Cumberland Plateau is perhaps the most floristically diverse physiographic 
province in Tennessee, as shown in a recent treatment of the woody plants of 
Tennessee (Wofford & Chester 2002). This treatment exemplifies that the 
Cumberland Plateau has both more native genera and native species (and lesser 
taxa) than any other physiographic province in the state and surprisingly a 
few more than the Blue Ridge which includes the Tennessee portion of the Great 
Smoky Mountains National Park and numerous other high mountain peaks. 

The highly dissected northern portion of the Cumberland Plateau in Ten- 
nessee (Fentress, Morgan, Scott, and Pickett counties) and Kentucky (McCreary 
Co.) has a unique habitat commonly referred to as rockhouses. These are am- 
phitheater-like semicircular recessions under overhangs and ledges of Penn- 
sylvanian sandstone. In addition to providing moist, sandy soil, other physical 
features unique to this habitat include lower light quality, lower summer and 
higher winter temperatures, higher relative humidity, and lower evaporation 
rates. Rare and/or unique taxa occur here at or behind the drip line and are 
only rarely in direct sunlight or rainfall. The flowering plant taxa of interest 
often have paler, more delicate leaves that probably would not withstand direct 
sunlight or heavy rainfall [see Walck et al. (1986) for an excellent review of sand- 
stone rockhouses with emphasis on ecology and evolution of endemic taxal. 

Several taxa closely related (with an asterisk) to Tennessee rockhouse 
endemics are often common in adjacent forests or outcrops. Examples of these 
sister species pairs are: Thalictrumclavatum DC.* and T. mirable Small, Arenaria 
glabra Michx.* and A. cumberlandensis Wofford & Kral, Ageratina altissima 
(L.) King & HE. Robins.* and A. luciae-brauniae (Fern.) King & HE. Robins., 
Silene viginica L* and S. rotundifolia Nutt., and Heuchera villosa Michx.* and 
H. parviflora Bartl. In addition, three pteridophyte taxa endemic to sandstone 
rockhouses in the eastern U.S. occur on both the Cumberland Plateau and Blue 
Ridge provinces of Tennessee (Walck 1986); these are: Vittaria appalachiana 
Mickel & Farrar, Trichomanes intricatum Farrar, and Trichomanes boschianum 
Sturm. The only rockhouse endemic that is also a Tennessee endemic is the 
herein described new species of Stenanthium. 


SPECIES DESCRIPTION 


Field observations for numerous seasons from Cumberland Plateau rockhouses 
have resulted in the discovery of yet another undescribed rockhouse endemic. 


450 BRIT.ORG/SIDA 22(1) 


This new taxon is most closely related to S.gramineum but the distinct differ- 

ences in morphology, restricted distribution, phenology, associated taxa, and 

habitat requirements warrant recognition and description of a new species of 

Stenanthium, to wit: 

Stenanthium diffusum Wofford, sp. nov. (Figs. 1, 2, 4). Tyee: TENNESSEE. Pickett Co. 
Pickett State Park, beneath and at the margin of rockhouse ledges and overhangs on Ladder 
Trail at crossing of Thompson's Creek, 36°33'18"N, 84°47'55"W, elev. 1597 {t, 23 Sep 2005, BE. 
Wofford 20051, with D.K. Smith, G. Bresowar, & S. Huskins (HOLOTYPE: TENN; ISOTYPES: BRIT, 
GH, MO, NCU, NY, US) 


St hiogrami {fini | differt intlorescentia m; aay floril i ly i ramis 

later ali ae i yniciula ca. ee cel dis spos yeITIS 34¢ in | : C 3-4 mm longis portatis, 
et folio cauli i inus q 15cm supra es 

Plants glabrous perennial herbs witha single stem up to | m tall, mostly shorter, 


arising from a shallow seated tunicate bulb 2-8 cm deep; the bulb 2 cm high, 1 
cm wide on reproductive individuals and surrounded by a dark, fibrous net- 
work of remains of previous years’ leaf bases. Roots fibrous, numerous, up to | 
mm wide. Leaves 7-10, linear, entire, up to 3 dm long, 1-3(-4) cm wide, apex 
acute to slightly rounded, sheathing, predominately basal, only | or 2 above the 
basal rosette, conduplicate basally, reflexed and flattened distally, lowermost 
leaves often reclining; basal and cauline leaves (non-bracteal leaves) 15 cm or 
less above ground level, bracteal leaves becoming progressively reduced toward 
the inflorescence axes. Inflorescence a terminal, bracteate, diffuse panicle up to 
3.dm wide of nodding, simple or compound racemes, the terminal portion race- 
mose, erect, and upto 1.5dm long or mostly shorter or absent, at least not longer 
than the lateral racemes; lateral branches 6-10, up to 15 in larger plants, each 
subtended by a bracteal leaf that becomes smaller toward the stem apex, axis 
of lateral branches up to 1.5 dm long. Flowers numerous, mostly bisexual or 
with the lowermost functionally staminate and the uppermost functionally 
pistillate, distinctly pedicellate, those on the mid-portion of panicle branches 
(5-)8-15(-18) mm apart and on slender pedicels (_-)1.6-4(-5) mm long, those of 
the terminal raceme, if present, of similar length; ultimate pedicel bracts 13 
mm long, pale margined, clasping at the base. Tepals 6, spreading to slightly 
campanulate, scarcely connate basally, imbricate, narrowly lanceolate, apex 
acute to acuminate, tortuous, white at anthesis with a pale green midrib, be- 
coming green and white margined with fruit development, persistent; sepals 
4.5mm long, 1.2 mm wide; petals 5 mm long, | mm wide, slightly narrower and 
longer than the sepals, scarcely tumid along the basal margins but not glandular 
or nectariferous. Stamens 6, in two series, antitepalous, scarcely epitepalous; fila- 
ments white, 0.7 mm long, recurved; anthers L-locular, yellow, 0.4 mm long and 
wide, reniform, basifixed, extrorse; dehiscing intoa peltate, dumbbell shaped disc; 
pollen yellow, ellipsoid. Gynoecium 3-carpellate, syncarpous, placentation axile; 
ovary about one-third inferior, white, 1 mm long, becoming green and glaucous 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 451 


Fic 1 uel typ Lee ah? ALL. (We ££, {INNCA , TENN) 


452 BRIT.ORG/SIDA 22(1) 


with fruit development, ovules up to 12, most aborting; styles 3, recurved, 0.4 mm 
long, persistent. Fruit a glaucous capsule 7-9 mm long, 3-4 mm wide; styles per- 
sistent and recurved: dehiscence septicidal; seeds fusiform, sometimes flattened 
on one side due to compression, 1-6/locule, 3-3.5 mm long, | mm wide, brown, 
with irregular wrinkles, flattened and slightly arcuate at the tip. 

Habitat, distribution, and phenology.—Stenanthium diffusum is restricted 
to sandy, moist soil on ledges and below rockhouse overhangs on the northern 
Cumberland Plateau of Tennessee (Fig. 5). It is presently documented from only 
five populations in Morgan, Fentress, Pickett, and Scott counties (Fig. 6). The 
largest are found in Pickett State Park where they are associated with rockhouse 
endemics Silene rotundifolia, Heuchera parviflora, Vittaria appalachiana and 
the federally endangered Arenaria cumberlandensis. lt flowers from mid-Sep- 
tember to early October with fruits maturing in mid-November. 


Additional collections examined. TENNESSEE. Fentress Co.:ca.6 mi S Pickett Park, shi eras 
ledges, sandstone, 6 Sep 1978, R. Kral 62661 (MO & VDB, as photocopy!); Twin Arches, 13 Oct _D. 
Eagar s.n.(VDB, as photocopy!)*. Morgan Co.: beneath sandstone ledges at crossing of Tennessee en 
ley Authority transmission line above tributary to Clear Fork River, 36°18'59'N, 84°46'46'W, elev. ca. 
1300 ft, 3 Oct 2005, B.E. Wofford 20053, with D.K. Smith & G. Sexton (TENN). Pickett Co.: acid sandy 
oak-pine woods above Thompsons Creek ca. | mi N Park Headquarters, 7 Jul 1973, R. Kral 50563; in 


SISNINT: p] 


sand of rockhouse, | lidden Passage Pickett State Park, IBA ug 1973, A. Sharp sn C1 ENN): Shai p Place 


Quad., Pickett State Park, Ladder Trail, series of sandstone rockhouses near Hwy. 154, in damp soil ol 


sandstone rockhouse overhangs, with hemlock, near Arenaria cumberlandensis, 07 Aug 1993, Milo 
Pyne 93-204(TENN, VDB, as photocopy!); moist sanc 
Ladders Trail, Pickett Park, 22 Sep 1974, R. Kral cece VDB, as photocopy); Pickett State Park, 
rockhouse on Hidden Passage Trail, ca. 100 yards E of its jet. with Group Camp Trail, 36°33'35'N, 
84°47'26'W, elev. 1538 ft, 26 Oct 2005, B.E. Wofford ce with G. Sexton, D. Estes, & J. Beck (TENN). 
Scott Co.: in moist, sandy soil at base of sandstone bluffs at Twin Arches, ca. 0.2 mi. w of Charit 
Creek. 36°32°30"N, 84°44'09"W. 26 Oct 2005, BE. Wofford 20052, with D.K. Smith, G. Bresowar, & S. 


Huskins (1 ane *The correct county for this specimen is Scott Co. 


stone ledges in hemlock-white pine forest above 


DISCUSSION 


Stenanthium diffusum is easily distinguished from S. gramineum by morphol- 
ogy, geography, phenology, and habitat (Table 1). Morphologically, the most ap- 
parent difference between these two taxa is the nature of the inflorescence. The 
inflorescence of S. diffusum is broader, up to 3 dm wide, more diffuse, and the 
terminal, wand-like raceme is absent or nearly absent (Fig. 2). The diffuse na- 
ture of the inflorescence of S. diffusum is due to the more numerous and longer 
secondary panicle branches that usually exceed 3cm in length. In S.gramineum 
the inflorescence is less diffuse and rarely exceeds 1.5 dm in width except in 
some individuals perhaps referable to var. robustum; in addition, the terminal 
raceme is always present and may comprise up to 20-70% of the total inflores- 
cence length (Fig. 3). Furthermore, in S. gramineum, secondary branches are 
infrequent and generally less than 2 cm long. 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 453 


Fic. 2. Stenanthium diffusum habit at anthesis. Image taken at type locality, Ladder Trail, Pickett State Park, Pickett 
County, Tennessee. 


454 BRIT.ORG/SIDA 22(1) 


Taste 1.Morphological and other comparisons of Stenanthium diffusum and S.gramineum. Quanti- 
tative values are based on measurements from specimens deposited in the University of Tennes- 
see Herbarium (TENN). 


S. diffusum S.gramineum 
Pedicel length at mid-portion of lateral (1-)1.6-4(-5) (0-)0.3-1.1(-1.7) 
inflorescence branches (mm) 
Distance between pedicels at mid-portion (5—)8-15(-18) (1-)3-7(-8.5) 
of lateral inflorescence branches (mm) 
Distance from ground to uppermost (3-)4-14(-15) (15-)22-66(-76) 
cauline leaf (cm) 
Distribution northern Cumberland 
Plateau of TN FL to TX,n to IN and PA 
Flowering period (at ca.same latitude) mid a to mid-Oct May to late Aug 
abitat moist, shady,sandstone — dry to mesic woodlands 
rockhouses and open areas 


Stenanthium diffusum and S.gramineum also may be separated by quanti- 
tative morphology, phenology, distribution, and habitat (Table 1). The flowers 
of the mid-portion of the lateral inflorescence branches of S. diffusum are sepa- 
rated from each other by =8 mm and are borne on distinct pedicels (=1.6 mm 
long) that often equal or exceed the subtending bracts. In comparison, the [low- 
ers of the lateral inflorescence branches of S.gramineumare sessile or subsessile 
on pedicels (<1.1 mm long) that are equal to or shorter than the subtending 
bracts and are separated from each other by <7 mm, giving the inflorescence a 
less diffuse appearance. Interestingly, although the pedicels on the lateral 
branches are sessile or subsessile, those of the terminal wand-like raceme are 
distinctly pedicellate and may be up to 5 mm long. Another reliable character- 
istic that readily separates S. diffusum and S. gramineum is the distance from 
the ground to the node of the uppermost cauline leaf (not bracteal inflores- 
cence leaf). The distal-most cauline leaf of S.diffusum is <14 cm above ground 
level whereas the distal-most cauline leaf of S. gramineum is >22 cm above 
ground level. Although somewhat difficult to quantify from herbarium mate- 
rial, there also appears to be a difference in leaf length/width ratio between 
these taxa, with the tendency toward a lower leaf length/width ratio in S. 
diffusum. 

Other non-morphological differences include: (1) phenological differences, 
ie., S.diffusum flowers from mid-September to mid-October and S.gramineum, 
at this approximate latitude, flowers from May to late August, (2) S. diffusum is 
a narrow endemic to the northern Cumberland Plateau of Tennessee in sand- 
stone rockhouse habitats characterized by siliceous soils, low light intensity, 
high relative humidity, lower evaporation rates, and lower summer and higher 
winter temperatures than the surrounding forests. The more widespread S. 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 


Fic.3. Stenanthium gramineum habit at anthesis. 


gt 


fl 


BRIT.ORG/SIDA 22(1) 


h, from holotype 


Fic. 4. Stenanthium diffusum, image of flowers at mid- 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 457 


Fic. 5. Habitat of Stenanthium diffusum. Sandstone rockhouse formations at type locality, Ladder Trail, Pickett State 
Park, Pickett County, Tennessee. 


gramineum occurs in a much broader range of ecological conditions in numer- 
ous habitats including rich woods, moist to dry woods, meadows, flood plains, 
prairies, and grassy balds. The following key is provided for ease of identitica- 
tion: 


1. Inflorescence a diffuse panicle up to 3 dm wide, the terminal racemose portion 
reduced or absent;flowers on mid-portion of lateral branches with distinct pedicels 
1.6-4 mm long and 8-15 mm apart; uppermost non-bracteal stem leaf 4-14 cm 
above ground level S. diffusum 
Inflorescence not diffuse, typically up to 1.5 dm wide, the terminal racemose por- 
tion present and up to 3 dm long; flowers on mid-portion of lateral branches with 
pedicels 0.3-1.1 mm long and averaging 3-7 mm apart; uppermost non-bracteal 
stem leaf 22-66 cm above ground level S.gramineum (including 


var. micranthum and var. robustum) 
CONSERVATION CONCERNS 


The northern Cumberland Plateau of Tennessee and adjacent Kentucky har- 
bors a suite of species commonly referred to as rockhouse plants. This unique 
flora is restricted to sandstone floors, ledges, and overhangs. There are several 


458 BRIT.ORG/SIDA 22(1) 


ad 
em 
aul 


an 
eee @epmeemen eae 


ar 


> oa] 
VA 
NC 
sc 
Legend 
\ ® Populations 
\ 
\ 100 
‘ TT Kilometers 


Fic 6 r + J Ly + Es | - £ Cy oy AT; 


critical factors required, in concert, to maintain these unique habitats: moist 
sandy soil, low light intensity, high relative humidity, low evaporation rate, and 
ower summer and higher winter temperatures than adjacent forests (Wofford 
& Smith 1980). 

The environmental factor most likely to have an immediate and negative 
impact would be removal or severe thinning of adjacent woodlands. This would 
increase light intensity and temperature while reducing relative humidity and 
moisture. An aggressive management plan should be placed into motion on 
public lands (especially at Pickett State Park and Forest and the Big South Fork 
National River and Recreation Area) aimed at protecting a part of this critical 
habitat and its associated flora. In addition to protecting a unique part of 
Tennessee's natural heritage, protection would provide opportunity for study 
of biological/evolutionary problems yet to be resolved, i. e., speciation and sis- 
ter pair relationships, micro and macroclimates, endemism, rarity, etc. There- 
fore, itis highly recommended that the Tennessee Department of Environment 
and Conservation and the U.S. Fish and Wildlife Service, in coordination with 
public land managers, conservation groups, and interested individuals, propose 


— 


_ 


WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 459 


both state and federal endangered status for Stenanthium diffusum. Hopefully, 
these efforts can, in concert, accomplish the necessary management and mis- 
sion plans to preserve this unique habitat and flora. 


ACKNOWLEDGMENTS 


I thank the directors/curators of MO and BRIT/VDB for providing images of 
specimens. Special thanks goes to the ey and graduate students at the Uni- 
versity of T for their « numerous field trips; to Dwayne 
Estes who carefully reviewed early drafts of the manuscript; to Guy Nesom who 
provided the Latin diagnosis; to Chris Fleming for constructing the distribu- 
tion map; to anonymous reviewers for their helpful comments, to Jerry Drown 
for providing the image of Stenanthium gramineum, and to the Hesler Fund, 
University of Tennessee, for funding field trips. 


REFERENCES 


FerNALD, M.L. 1946. Stenanthium in the eastern United States. Rhodora 48:148-152. 

Gates, R.R.1918.A systematic study of the North American Melanthiaceae from a genetic 
standpoint. J. Linn. Soc. Bot. 44:131-172. 

JOHNSON, R.G. 1969. A taxonomic and floristic study of the Liliaceae and allied families in 
the southeastern United States. Ph.D. dissertation. West Virginia University. 

USDA, NRCS. 2005.The PLANTS Database, Version 3.5 (http://plants.usda.gov). Data com- 
piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge, 
LA. 70874-4490 USA. 

Utec, F.H.2002. Stenanthium.|n:Flora of North America Editorial Committee. Flora of North 
America north of Mexico. Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford Univ. 
Press, New York, New York. Vol. 26. Pp. 88-89. 

Voss, E.G. 1972-1996. Michigan flora, 3 parts. Part 1,Gymnosperms and monocots. Bloom- 
field Hills and Ann Arbor.P.418 

Watck, J.L., JM. Baskin, and C.C. Baskin. 1986. Sandstone rock houses of the Eastern United 
States, with particular reference to the ecology and evolution of the endemic plant 
taxa. Bot. Rev. 62:311-362 

Worrord, B.E.and R.Krat.1979.A new Arenaria from the Cumberlands of Tennessee. Brittonia 
31:257-260. 

Worrorb, B.E. and E.W. CHester. 2002. Guide to the trees, shrubs, and woody vines of Ten- 
nessee. The University of Tennessee Press, Knoxville. 

Worrorb, B.E. and D.K. SmitH. 1980. Status report on Arenaria cumberlandensis Wofford & 
Kral. Report submitted to U.S. Fish & Wildlife Service, Atlanta. 

Zomterer, W.B. and W.S. Jupp. 2002. Resurrection of segregates of the polyphyletic genus 
Zigadenus 5.l. (Liliales: Melanthiaceae) and resulting new combinations. Novon 12: 


460 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


RiA Loonuizen. 2005. The Elder in History, Myth and Cookery. (ISBN 1-903018- 
31-5, pbk.). Prospect Books, Allaleigh House, Blackawton, Totnes, Devon TQ9 
7DL, United Kingdom. (Orders: The David Brown Book Co., PO. Box 51], 
28 Main St., Oakville, CT 06779, U.S.A., 860-945-9329, 860-945-9468 fax, 
Email: david.brown.bk.co.snet.net). $17.95, 140 pp., 51/2" x 71/4" 


This petite book is an affectionate ode to the European elder (Sambucus nigra). It includes chapters 


on elder botany, mythology, and cuisine, as well as medicinal and other uses for the plant. Ina chap: 


of 


ter titled “The Healing E Ider,” the author reviews traditional medicinal uses of the elder. The proper 


harvest and drying of eldenlowst>: e uSeHOET IES and leaves is ite Piesnone io making. oint- 
} 


ments, syrups pouivs es 


that the E | tiviral and ad ic eff pporting i liti easa treat- 
ment for ee A chapter named “The C acme! “Ider” describes Bidens recipes, which sneha e such 


delicacies as elderf ia tuack ureter wine Lula jelly, CEA sour soup, and recipes for 


the Jew’s Ear mushroom (Auric ie-judae), which grow r wood. These reci- 
J 


pes can be prepared using the American elder na us ioe nsis). However, caution should be 
can | rt ff t. Pep 


used because large amounts of elderberries and bar red t ee the 


book are illustrations, photographs, and quotes ae to the elder. The author es vith a quote 

by the poet Seamus Heany: “I love its blooms like saucers brimmed with meal, its berries a pine of 
shot, a buoyant spawn, a light bruised out of purple. Elderberry? Itis shires dreaming wine.’—Marissa 
N. Oppel, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A. 


SIDA 22(1): 468. 2006 


A NEW SPECIES OF DODECATHEON (PRIMULACEAE) 
FROM THE NORTHERN COAST RANGE OF 
OREGON AND WASHINGTON 


Kenton L. Chambers 


Department of Botany and Plant Pathology 
Oregon State University 
2082 Cordley Hal 
Corvallis, Oregon 97331-2902, U.S.A. 


ABSTRACT 


Dodecatheon austrofrigidum is described as a species occupying scattered sites in the Coast Ranges 
of northwestern Oregon and adjacent W, ere USA. It is similar in floral morphology to, but 
widely disjunct from, D. frigidum, a species of the interior mountains and arctic tundra of Alaska 


and northern British Columbia. It also is ecologic cally distinct, being associated with streams, water- 
falls, cliffs, rocky river banks, and high elevation moist grassland and basalt talus slopes in a mari- 
time climatic zone. Its relationships are with members of Sect. Dodecatheon, but its seedling mor- 
phology is different from any described thus far in the genus. Morphology of the seed testa is 
illustrated by scanning electron microscopy. 


RESUMEN 


Dodecatheon austrofrigidum se describe como una espec upa lugares espaciados de la 
dillera Costera del Noroeste de Oregon y de W. ashington, U SA Es en en 2 puorteloe al i 


pero muy disyunta de D.frigidum le Alaska 


y norte de British Colum bia. imapien es ecologicamente diferente, estando asociada a arroyos, 
cascadas, acantilados, lechos rocosos de rios, y praderas humedas a gran altitud y laderas basalticas 


n una zona climatica marina. Esta relacionada con los miembros de la Sect. Dodecatheon, pero la 
morfologia de sus plantula liferente de cualquiera de las hasta gee descritas en el género. Se 


Ha 


ilustra la moriolose de testa seminal al microscopio electrénico de barr 


INTRODUCTION 


The northern Coast Ranges of Oregon and immediately adjacent Washington, 
especially on the higher peaks, are known to harbor a small suite of endemic 
plant species. Prominent in this list are Cardamine pattersonii LF Hend., 
Castilleja chambersii M. Eggers & R. Meinke, Erythronium elegans P Hammond 
& K.L. Chambers, Filipendula occidentalis (Wats.) Howell, and Saxifraga 
hitchcockiana Elvander. A floristic affinity to the Olympic Mountains and 
coastal Alaska is suggested by more northern taxa whose ranges extend south 
to this region. These include Carex macrochaeta C.A. Mey., Cladothamnus 
pyroliflorus Bong., Erigeron peregrinus (Pursh) Greene var. peregrinus, Lewisia 
columbiana var. rupicola (English) CL. Hitchc., Prenanthes alata (Hook.) Dietr, 
Saxifraga caespitosa var. emarginata (Small) Rosend., Senecio flettii Wieg.. and 


SIDA 22(1): 461 — 467. 2006 


pas 


462 BRIT.ORG/SIDA 22(1) 


Synthyris schizantha Piper (Chambers 1973, 1974). Saddle Mountain, Clatsop 
County, Oregon, is a well known botanical site where many of these species 
have been found (Detling 1954). 

Also known from Saddle Mountain, near the 1000m summit, is a dwarfed 
form of Dodecatheon, which the monographer, H,J. Thompson (1953, p. 118) re- 
ferred, with some hesitation, to D. pulchellum (Raf.) Merrill (his D. radicatum 
Greene). A number of years ago! was directed to populations of vigorously grow- 
ing examples of this plant along the Trask River, Tillamook County, Oregon, 
which I brought into cultivation. This tentative new species was then made 
known to interested botanists under the name Dodecatheon austrofrigidum, 
which has now come into general use for the plant. [here provide a description 
and further documentation of this species. 


Dodecatheon austrofrigidum K.L. Chambers, sp. nov. (Figs. 1, 2) Type: US.A. Or- 
EGON. Tillamook Co. Trask River Rd., 6.1 mi E of junction with Long Prairie Rd., TIS, R8W, 
§28, elev. ca. 200 ft, in patches of moss on bare, cea: rocky bank of the ees River, be- 


tween low and high water marks, with Saxif llii, S. mertensiana, Mi 
13 May 1989, K.L Chambers ae OTYPE: OSC: ISOTYPES: FSU, G, MARY, MO, NY, OSC, UC, 
US, WS, WTU 


a 


Planta ad Dodecatheon frigidum et Y aes — Sea variis ase ede ci seis glabro dentibus 
triangularibus acutis lobis corollae ibe 


] 


eris connectivis 
antherarum purpureis laevibus stigmate non capitato ab D. fr an radicibus numerosis fibrosis 
validis caudice plerumque per annos paucos persistenti differt, ab D. dentato corolla lavandula non 
alba differt; chromosomatum numerus 2n = 88 


Perennial herb; roots numerous, stout, fibrous, often producing adventitious 
buds; caudex usually short-lived, with subsequent years’ vegetative growth aris- 
ing from a terminal bud or from small buds on the roots; leaves in a basal ro- 
sette, glabrous, variable, 2.5-30 cm long, 0.7-7 cm wide, ovate to broadly or nar- 
rowly elliptic, gradually or abruptly tapering to the petiole, margins entire to 
irregularly sinuate-dentate or denticulate; flowering stems scapose, 5.5-45 cm 

igh; inflorescence a bracteate umbel of 1-7 pedicelled flowers, bracts 2.5-10 
mim long, pedicels 4-55 mm long, glabrous to glandular-puberulent, especially 
distally; ealyx 5-11 mm, tube 1-2.5 mm, lobes 3-9 mm, triangular, acute; corolla 
lobes 9-23 mm long, 2-6 mm wide, lavender, tube 1-2 mm long, white with a 
sinuous purple line where reflexed; anthers purple, +.5-8 mm long, filaments 
0.5-1 mm, free or slightly united, purple, connective smooth, purple; style ex- 
serted, stigma not capitate; capsule 6-16 mm long, 3.5-5.5(-7) mm wide, teeth 
ca. 1/5 of total length, dehiscence operculate or valvate; seeds 1-2 mm, irregu- 
larly rounded or prismatic, foveolate (Fig. 2); seedlings with petiolate cotyle- 
dons, the first principal adult root produced adventitiously from the apex of 
the hypocotyl, the first true leaf petiolate, arising between the cotyledons (Fig. 

. The chromosome number is tetraploid on the base of x = 22 characteristic of 
the genus. 


CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 463 


Fic. 1.Dodecath trofrigidum K.L. Chambers. A. Habit. Scale=2 cm. B. Flower. Scale=7 mm.C. Capsule with opercu- 


lum. Scale=7 mm. 


464 BRIT.ORG/SIDA 22(1) 


Fic. 2. Seed. Scale=0.1 mm. 


Distribution.-Ranging from 30-1200 m, at scattered sites from the south- 
ern Olympic Mountains, WA (Mt. Colonel Bob) to Tillamook County, OR, both 
at high elevations in moist, grassy turf (Saddle Mountain and Onion Peak, 
Clatsop Co., OR) and in forested regions, where the habitats include moist ba- 
saltic slopes, talus, and cliff-faces, especially near waterfalls and streamlets. 
Further populations occur at low elevations in Tillamook Co.,on bare or m Ossy 
basalt rock banks of rivers, between the low- and high-water marks, in the win- 
ter flood-scour zone (Raven 1995, 1996). Plants in this peculiar habitat probably 
become established by seeds washed down from the mountain populations, and 
their tenacious mass of strong roots, penetrating crevices in the rock, anchors 
them from being washed away during the period of winter flooding. The spe- 
cies is allopatric in range to all other members of the genus. 

Flowering. —April-July, depending on habitat and elevation. Flowers are 
self-fertile and capable of self-pollination in cultivation. 

Relationship.—Thompson (1953) recognized three sections within the ge- 
nus, differentiated by characteristics of the stigmas, capsules, seeds, and seed- 
lings. The new species is best assigned to Sect. Dodecatheon, with the proviso 
that D. frigidum Cham. & Schltdl. should also be placed in this section, rather 
than Sect. Purpureo-tubulosa of Thompson. The corrected placement is strongly 
supported by molecular evidence provided by Mast et al. (2004) and by the 


— 


CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 465 


marked floral similarity between D.frigidum and D. austrofrigidum. Mast et al. 
(2004) show a strict consensus tree based on seven cpDNA regions, in which D. 
frigidum is sister to the group of other taxa in Sect. Dodecatheon, rather than 
forming a clade with D. conjugens Greene in Sect. Purpureo-tubulosa, as pro- 
posed by Thompson. In this DNA-based phylogeny, D. austrofrigidum is paired 
in Sect. Dodecatheon with D. dentatum Hook. ssp. utahense (N. Holmgren) 
Kartesz (perhaps itself a distinct species). 

Dodecatheon austrofrigidum differs in two significant features from the 
morphological characterization of Sect. Dodecatheon given by Thompson 
(1953). These, and an improved knowledge of the seed testa based on SEM stud- 
ies (Fig. 2), will be discussed in turn. Most important, I believe, is that seedling 
morphology and development are distinctly different than had been described 
for this section. In D. austrofrigidum, the seed begins with two petiolate cotyle- 
dons (as Thompson, 1953, Fig. lg, illustrates for D. pulchellum). The first true 
leaf, itself petiolate, next arises from between these cotyledons. Finally, a strong, 
major adult root forms adventitiously at the base of this first true leaf, between 
the cotyledons, replacing the seedling’s initial but ephemeral root system (Fig, 
3). In the seedling of D. pulchellum (D. radicatum) illustrated by Thompson 
(loc. cit.), the first true leaf is linear-spatulate, not petiolate, and arises adventi- 
tiously from the hypocotyl well below the cotyledons; the new adventitious 
major root grows downward from that same point on the hypocotyl. Neither of 
the other two seedling types illustrated by Thompson (loc. cit.) resembles that 
of D. austrofrigidum. 

The mature capsules of D. austrofrigidum differ in their dehiscence mode 
from that described for Sect. Dodecatheon (Thompson, op. cit., p. 79). The sec- 
tion is characterized as having valvate capsules, but in D. dustrofrigidum, they 
are most often operculate instead. Capsules are thin-walled below and have an 
indurate apex where the cells are thick-walled and isodiametric. As these cells 
dry at maturity and shrink, tensions develop that rupture the tissue and open 
the capsule for seed dispersal (Fig. 1C). A transverse rupture through the tip 
region causes an operculum to free itself, carrying with it the dried style. Usu- 
ally five lengthwise splits also occur at the same time, but some of the five teeth 
thus formed may in turn split in half, so that dehisced capsules may be found 
with up to ten teeth. In a sample of 181 capsules, the average number of teeth 
was 6.28, the numbers of capsules with 5, 6, and 7 teeth being approximately 
equal. The initial five splits can often be observed in the operculum (Fig. 1C). 
Valvate dehiscence, where the lengthwise splits reach the apex, is usually noted 
in smaller, less robust capsules. 

Seeds of D. austrofrigidum (Fig. 2) conform to the description of Sect. 
Dodecatheon (Thompson op. cit., p.75-76). They are irregular in shape but of - 
ten flattened on two or more sides, becoming even more prismatic than in 
Thompson's drawing (op. cit., Fig. lf). They do not resemble the membranous- 


466 BRIT.ORG/SIDA 22(1) 


t 


¢.3. Seedling, showing petiol yled 1 first leaf. A major adventiti tight, repl he initial, short- 
lived seedling hypocotyl and root at the left. Scale=5 mm. 


margined seeds of Sect. Capitulum (Thompson loc. cit.). The testa is formed ini- 
tially of tightly packed bulbous cells, which at maturity deflate into irregular, 
rimmed, dish-shaped foveae (Fig. 2), best visualized by SEM. In view of these 
newly published observations on D. austrofrigidum, it is likely that a survey of 


CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 467 


seed, seedling, and capsular morphology in all the species will contribute toan 
improved understanding of intrageneric relationships. 


Cited collections: U.S.A. OREGON. Clatsop Co.: Onion Peak, elev. 3064 ft, T4N, RIOW, $22, 29 Jun 
1971, aus a 3169 (CAS, OSC); same, 21 Jul 1971, Chambers 3258 (OSC); Saddle Mtn, T6N, R8W, 
$33, 29 972, D. Jaques 973 (OSC); same, elev. 2600 ft, TON, R8W, S19, 24 May 1950, Detling 6550 
(CAS, ae "v0 madals Mi elev. 3200 ft, 10 Jun 1928, G. R Patterson s.n. (ORE). Tillamook Co.: Trask 
River Rd.a ,on pasa pani ef Trask R. ieee 1980, Chambers 4694 (OSC); same, 10 
Jun 1979.6. Lewiss.n (OSC. S d.aboy k Ck., elev. ca. 3200 ft, TZN, R7W, S13 1, photo, 
Jun 1981, G. Lewis s.n.(OSC); near Diamond Falls, he State Forest, elev. 1480 ft, T2N, ROW, S32, 
13 Jun 2001, Chambers 6300 (OSC), Kilchis Falls, Tillamook State Forest, elev. 1540 ft, TIN, R8W, $3, 
13 Jun 2001, Chambers 6299 (OSC). WASHINGTON. Grays Harbor Co.: Mt Colonel Bob, above Moon- 
shine =e elev. 3800 ft, T23N, R8W, S18, 20 Jul 1983, E. Alverson 590 (OSC); same, 6 Jul 1994, D. Davis 

cific Co.: Willapa Hills, headwaters of West Fork of Grays R., elev. 1500 ft, TILN, R7W, 
S10, : yee eee Powell 1324 (OSC): same, elev. 1800 ft, 23 Apr 1995, J. Powell 1410 (OSC). 


ACKNOWLEDGMENTS 


The author gratefully acknowledges the help of the following colleagues in this 
study: Dave Buchholz, Dan Clough, Thea Cook, Aaron Liston, Cathy Maxwell, 
Barbara Moore, Andrea Raven, James Reveal, Larry Scofield, Rena Schlachter, 
Clint Smith, and Al Soeldner. John Megahan prepared the illustrations. Special 
thanks are extended to Gerald Carr, who made the cytological preparations, 
and George Lewis, who first directed me to populations of Dodecatheon 
austrofrigidum on the Trask River. Austin Mast and Harry Thompson provided 
prompt and helpful reviews. 


REFERENCES 


CHAMBERS, K.L. 1973. Floristic relationships of Onion Peak with Saddle Mountain, Clatsop 
County, Oregon. Madrono 22:105-114. 

CHameers, K.L. 1974. Notes on the flora of Clatsop Co., Oregon. Madrono 22:278-279. 

Detune, L.E. 1954. Significant features of the flora of Saddle Mountain, Clatsop County, 
Oregon. Northwest Sci. 28:52-60. 

Mast, A.R., D.M.S. Feuer, S. Ketso, and E. Conti. 2004. Buzz-pollinated Dodecatheon originated 
from within the heterostylous Primula subgenus Auriculastrum (Primulaceae):a seven- 
region coDNA phylogeny and its implications for floral evolution. Amer. J. Bot. 91: 
926-942 

Raven, A.N. 1995. Dodecatheon austrofrigidum monitoring on the Trask River Peninsula, 
Tillamook County, Oregon. Berry Botanic Garden, Portland, OR. 

Raven, A.N. 1996. Summary of six years of research on Dodecatheon austrofrigidum, the 
Frigid Shootingstar, at the Trask River Peninsula, Tillamook County, Oregon. Berry Botanic 
Garden, Portland, OR. 

THompson, H.J. 1953. The biosystematics of Dodecatheon. Contr. Dudley Herb. 4:73-154. 


468 BRIT.ORG/SIDA 22(1) 


Book REVIEW 
WILLIAM C. BurGer. 2006. Flowers: How they Changed the World. (SBN 1-59102- 
407-2, hbk.). Prometheus Books, 59 John Glenn Drive, Amherst, NY 14228- 
2197, U.S.A. (Orders: 716-691-0133, 716-691-0137 fax, http://www. 
prometheusbooks.com/). $23.00, 337 pp., b&w figures, color photographs, 
glossary, and notes, 5" X 7" 


William Burger has written a book that is hard to put down. Flowers: How they Changed the World is 


a fantastic introduction to the angiosperms, their evolutionar y idv ces, and the roles plat gh play in 


the evolutionary history of man. This book is written in such a way ah at it will appeal to those read- 

ers looking for an introduction to flowering plants. However, it also provides the seasoned readers 

much food for thought. The writing style is very ss and conversational with a good deal of wit, 
k 


yet includes an abundance of information. Th s many topics including: what is a flower, 


the purpose of flowers, flower friends, ne enemies, how to distinguish flowering plants, what 


makes flowering plants special. It also discusses how primates, people and flowering plants relate to 
rer, which 


each other and in the end how flowers changed the world. Topics flow well into each ot 
greatly adds to the readability of the text. 


The author is clever about ke ing new subjects and making these topics non-threatening. 


For exam ie the book starts with introductions to four plants, a geranium, rose and two lilies. The 
author hints at the importance of distinguishing traits between monocotyledons and dicotyledons, 


but doesn’t delve into more detail until later chapters; smartly building on these unassuming intro- 
ductions made in the first chapter. This book contains well-written discussions and a wonderful 
introduction to major evolutionary advances in plant reproductive biology. Among these are how 
pollen helped flowering plants do well on land, the value of deciduous leaves, the biology related to 
losing leaves from a plant, and the evolutionary advances in plant defenses of chemicals that help 
prevent peEovory and disease. The author includes many other interesting discussions on subjects 


such as the importance of out- sea in eee and how plants have adapted ways to 2 out- 


nrere 


bree ding, as well as the diversity in fo re of [lowerin ng pla ants. The i : c ay 5 


_ 


cover information on how plants have been oa tats to people, focusing on mankind's develo 
ment of agriculture and the importance of biodiversity to our lives and the Earth. There is ales. a 


section that summarizes the important role plants have played in creating our world and the modern 


lives of man in ten major stages. 
lowers: How they Changed the World by William Burger is an extraordinary and interesting 
kgrounds and inter- 


jon 


book to read; full of engaging information for readers of all ages, educational bac 
ests. The text is written in an easygoing style and covers a wide range of topics related to the evolu- 
tion and biology of flowering plants. The author has also gone above and beyond typical notes for the 


rea der; he eeiauceh both ee mation < | refer rences s for each chapte1 topic accented by his own edu- 


cated val Lucke ydoo, Herbarium, Botanical Research Institute of Texas, 509 Pe- 
can Street, Fort Worth, TX a 4060, U.S 


SIDA 22(1): 468. 2006 


THE LECTOTYPIFICATION AND 197" CENTURY HISTORY OF 
CROTON ALABAMENSIS (EUPHORBIACEAE SS.) 


Kenneth J.Wurdack 


Department of Botany 
Smithsonian Institution 
PO. Box 37012, NUNH MRC-166 
Washington, DC 20013-7012, U.S.A. 
wurdackk@si.edu 


ABSTRACT 


A detailed account is presented of the historical | he di of Croton alabamensis 
var. Ts one eco E.A. Smith in 1877 avd its subsea ent study by 19th century 


American botanists. A lectotype is designated and its 


RESUMEN 


Este articulo presenta una descripcion detallada de los eventos relacionados con el descubrimiento 
de Croton alabamensis var. alabamensis (Euphorbiaceae ss.) por E.A. Smith en 1877 y su estudio 
subsiguiente por botanicos en el siglo XIX. Ademas, se designa un lectotipo para esta especie y se 
discute su origen. 


INTRODUCTION 


The progress of botany in the southern United States during the latter half of 
the 19th century has generally not been critically studied. Most of the major 
figures have yet to be treated by more than brief biographical sketches. Histori- 
cal narrative is usually traced from the perspective of a particular person or 
institution, but a few rare southern plants (e.g., Croton alabamensis E.A. Sm. ex 
Chapm., Dionaea muscipula Ellis, Elliottia racemosa Muhl. ex EIL., Franklinia 
alatamaha Bartr. ex Marsh., Shortia galacifolia Torr. & A. Gray, and Torreya 
taxifolia Arn.) have attracted sustained botanical interest from discovery to the 
present day and are therefore of noteworthy historical interest. Croton 
alabamensis contains an especially rich and complete historical record that is 
chronicled here to recover lost details about the type collection and reveal ex- 
tensive and collegial interactions among the network of botanists of the day. 
Croton alabamensis is, in habit, the largest native United States representa- 
tive of the tropical-centered family Euphorbiaceae s.s. The species has a disjunct 
distribution between Alabama and Texas and contains two morphologically 
well-differentiated, geographically-named varieties (vars. alabamensis and 
texensis Ginzbarg, respectively). Croton alabamensis is readily distinguished 
from other North American species of Croton L. by its shrubby habit, petaliferous 
female flowers, and covering of lepidote scales that give the abaxial leaf sur- 


— 


SIDA 22(1): 469 - 483. 2006 


470 BRIT.ORG/SIDA 22(1) 


faces a silvery appearance. Webster initially suggested its affinities were with 
section Andrichnia Baill, but later referred it to section Lamprocroton Mull. Arg. 
and then section Argyrocroton (Mull. Arg.) G.L. Webster (Webster 1967, 1996, 
Webster et al. 1996). Recent molecular phylogenetic studies have affirmed a sis- 
ter relationship between the two varieties and suggested new affinities with a 
small “basal” Croton group (Moacroton clade) containing Cuban endemics 
Moacroton Croizat and Cubacroton Alain and other Croton species (mostly sec- 
tion Corylocroton G. L. Webster) ranging from the Caribbean to Central and 
South America (Berry et al. 2005; van Ee & Berry 2005, Wurdack et al. 2005; 
van Ee et al. in press). 

Croton alabamensis var. alabamensis is known from central Alabama and 
a single probably erroneous report from central Tennessee (see below; Farmer 
& Thomas 1969; Wofford & Chester 2002). In Alabama the typical variety oc- 
curs in Tuscaloosa County along the Black Warrior River and in adjacent Bibb 
County from the Cahaba River drainage. It was first discovered in Bibb County 
by Eugene A. Smith in 1877. Ironically, Smith was apparently unaware of the 
populations along the Warrior River in his home county, Tuscaloosa, which 
were found by Roland M. Harper in 1905 (Harper 1906b). Croton alabamensis 
var. texenesis was an unexpected discovery in 1989 from Texas, 1000 km away 
from the typical variety (Ginzbarg 1991, 1992; Aplet et al. 1994). 

The Alabama plants from the Black Warrior and Cahaba river systems are 
separated by ca. 40 air km but show morphological, cytological, genetic, and 
ecological differences (Farmer 1962; Farmer & Thomas 1969; van Ee et al., in 
press). The plants are semi-evergreen and undergo meiosis and bloom in early 
spring (February and March) from overwintering flower buds formed the prior 
season. In Bibb County the Croton forms dense understory thickets from 1-2.5 
m tall at 76-91 m of elevation in thin soils over limestone or dolomite above 
waterways within an 8 km radius of Pratt’s Ferry, an area roughly bounded by 
Schultz Creek on the west and Six Mile Creek to the east. The Cahaba River and 
its nearby tributaries cut steeply through the bedrock to form numerous dry 
rocky bluffs on which the Croton is especially abundant. 

Pratt’s Ferry has been reported to be the type locality of Croton alabamensis 
var. alabamensisand, as one of the most easily accessible localities for the plant, 
has been the source of numerous recollections. Pratt’s Ferry was a public ferry 
crossing of the Cahaba River just south of the community of River Bend and 
was probably named (although the Pratt family had many members in the area) 
for Mary Pratt (1800-1882), who ran the ferry from 1854 through at least the 
Civil War (Ellison 1984). Pratt’s Ferry and aptly named River Bend are ata large 
bend in the river where a ferry crossing could be established with relative ease. 
An iron bridge was built at the crossing in 1902 (Harper 1906a; the old roadbed 
is evident on the south side of the river and an iron pylon for this bridge re- 
mains on the north shore, pers. obs.) that was superseded by the present bridge 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 471 


crossing of County Road 26 just downstream from (west of) the original bridge. 
The exact nature of the ferry landings has been obscured. However, based on 
topography (ie., limitations imposed by adjacent bluffs), the ferry route appears 
to have diagonally spanned the river with the northern landing upstream and 
the southern landing downstream of the current bridge. In the vicinity of Pratt’s 
Ferry, the Croton presently occurs on the north side of the river ina small popu- 
lation immediately west of the current bridge and on the south side as a more 
or less continuous band east of the bridge for at least 1.6 km (pers. obs.). 

Despite a sustained interest in Croton alabamensis since its discovery in 
1877 and concomitant visits to Bibb County by many botanists, the flora in the 
area has only recently been documented and found to be rich in endemic and 
disjunct taxa (Allison & Stevens 2001). Most of the endemics occur on ecologi- 
cally sensitive patchy glades (Ketona Glades) developed over Ketona Dolomite. 
Populations of the Croton in the Cahaba River drainage are often found around, 
but not restricted to, the fringes of the Ketona Glades (pers. obs.). Despite occa- 
sional logging activities, the region appears relatively unspoiled, remote, and 
undeveloped today and similar toa vintage 19" century riverbank photograph 
(Davenport 1988:40). A very different picture ges when looking at the his- 
torical record of regional industrialization. Taken in that light, Charles Mohr’s 
dire prediction over 100 years ago that the survival of the Croton in its native 
habitat “will in all probability be of short duration” may not have been un- 
founded (Mohr 1889). The limited distribution of both Croton varieties, as well 
as the Ketona Glade flora, has caused conservation concerns. Some habitat in 
Bibb County has been recently preserved (ie., The Nature Conservancy's Bibb 
County Glades Preserve). 


Discovery of Croton alabamensis 

Eugene Allen Smith (1841-1927), professor of geology and mineralogy at the 
University of Alabama, was appointed State Geologist in 1873 of the newly re- 
established Alabama Geological and Agricultural Survey. He began surveying 
the geology of the state during summer expeditions in a custom-built field 
wagon (modeled after a horse-drawn ambulance and preserved at the Univer- 
sity of Alabama) in a manner documented in photographs (see Wolfe 1983:99) 
and in his copious geological field notes (Smith 1870-1889). Although the origi- 
nal field notebooks were lost, transcriptions made during a 1930's Depression- 
era public works project (probably WPA or preceding agencies) remain and al- 
low a detailed picture of Smith’s field activities to be reconstructed here. His 
recurring interest in Bibb County was economically oriented toward an inves- 
tigation of coal and iron deposits that had originally made the region a leader 
in antebellum iron production (Ellison 1984). In the mid-19" century the Bibb 
County countryside was dotted with ironworks, mill dams, stone quarries, 
mines for ore (limonite) and coal, and associated communities. Today few traces 


472 BRIT.ORG/SIDA 22(1) 


remain of this industrialization and the environmental degradation it caused 
is scarcely evident. Once thriving industrial communities are now extinct (ie., 
Tionis) or severely depopulated (i.e., River Bend and Six Mile). 

Smith first explored Bibb County in 1873 and in more detail in 1875. On 
August 17, 1875 he collected ore samples and explored associated Brighthope 
ironworks or “bloomery” (originally called Little Cahaba Furnace and in op- 
eration by 1850; Ellison 1984), the first blast furnace in the county and about 
140 m above a wooden dam (Browne’s Dam) on the Little Cahaba River that 
provided its power. The seats for the dam timbers appear today as a series of 
steps cut into a riverbank rock outcrop at the Bibb County Glades Preserve near 
Bulldog Bend (pers. observ; Ellison 1993:51). The apparent present day natural 
state of this area, which contains some of the finest Ketona Glades, is remark- 
able considering the past destructive activities engendered by the adjacent iron- 
works, although it is also possible that disturbance supported or enlarged the 
glade community. The following day (ie., August 18,1875) Smith continued from 
the Sinks (sec. 26, T24N, RIOE) on lower Six Mile Creek, where the creek flows 
underground except at flood stage, southwestward to Pratt’s Ferry, where he 
noted fossiliferous “limestone” containing the gastropod Maclurites magnus 
LeSueur. It seems improbable that the Croton escaped visual detection, even if 
not considered noteworthy at the time. 

The trip credited with the discovery of Croton alabamensis was from July 
18 to August 4, 1877. On July 18, Smith left Tuscaloosa and on July 24 his geo- 
logical party set up camp on the north side of the Cahaba River near Pratt’s 
Ferry. On the way, Smith described an area near Schultz Creek “where the Coal 
Measures are cut olf by a fault bringing up Knox Dolomite which ina series of 
glades or bare hills, with crumbling and much decomposed Dolomite stretches 
N.E. for a considerable distance” (entry for July 21; Smith 1870-1889). This is on 
the presently depauperate western end of the Ketona Glades zone and suggests 
that an extensive loss of glade habitat has since occurred. “Knox Dolomite” as 
used by Smith would have broadly included Ketona Dolomite which was named 
in 1910 (see Geological Survey of Alabama 1894) 

The label with Smith’s first collection of the Croton notes “Privet or Privy, 
Pratt’s Ferry. 26. 7. 77” (see Fig. 1). | interpret the date notation as July 26, 1877. 
Privet is probably derived from the vernacular for Eleagnus spp., which Croton 
alabamensis vegetatively closely resemble. Smith rarely recorded botanical 
observations in his geological field notes, and July 26 is no exception. On that 
date, Smith spent the morning west of Pratt’s Ferry on the north side of the 
river and then after lunch crossed over the river. On July 28, 1877, however, he 
makes a record of the Croton from a spot | estimate to be near the head of Little 
Schultz Creek, 3 km downstream from Pratt's Ferry. His field notes for that date 
read: 


_ 


{4 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 473 


2( 774058 ) 
a RS 
SOHAL HERS! 


; 
1UM OF CHARLES MOHR 


Presented ta the Smithsonian Institution, 1901 


terlom. Ulaba KemendS ty se 38 


xX 


STAT. EF HABITAT, 
ee -_ 
PML Fer 


HAE li are | iis 2 es : ee A Ce tra i 
TU eee ee 
Fic. 1. Crot, Lak G Dy (Pay ME aN EY gt i ra | * acclacedt hin Ila-ti fk pie ay soci- 
Il label in bottom left ( | li ) read “Type S Spm: pecan’ [hand of C. Mohr} "Prive or 
Privy. Pratts ay re iE i velonals) to Ala. Hoel "hand of E A. Smith] f Alak 
cant cate I y. Discovered by or E. ‘ Smith, August 27" 1879. — forms 


the banks of the little Cahaba ri bl ilM y” [hand of C. Mohr]. 


474 BRIT.ORG/SIDA 22(1) 


“Went with Mr, Mat Wallace in his skiff from opposite the point of the mountain [eastern end of Big 
Mountain] down nearly opposite the ore bank. The limestone bluffs furnished exactly the proper 
ground for the greatest variety of ferns. Amongst those I had not seen before were Adliantum|,. capil- 
lus-veneris, Cheilanthes alabamensis and one other which resembles it, but has lobed pinnules in- 
stead of entire ones. After climbing through an almost impenetrable thicket of what the natives call 
privet [Croton pasa and another shrub now in flower [probably Forestiera ligustrina Poir] 


and extremely fragran x thousands of bees we came to our ore bed” (Smith 1870-1889). 


Smith later recalled “the ie ocality of Croton alabamense, is Pratts Ferry in 
Bibb County, on [the] Cahaba River and upon limestone. Date of discovery, Au- 
gust 1877” (Smith 188ld; confirmed in Smith 1907). Smith probably encoun- 
tered the Crotonat other times during this 1877 trip, and assuredly so (based on 
present distribution) on August | while on the Cahaba River bluffs, 0.4 km north 
of where Cottingham Creek joins the river, while examining where “Knox Do- 
lomite” crosses. Smith appears not to have revisited the Pratt’s Ferry area until 
August 1889, based on field note chronology and referring to his “observations 
of 12 years ago” (ie., 1877). Mohr variously credits the discovery to 1874 (Mohr 
1898, 1901), 1877 (Mohr 1889), or 1879 (Mohr 1887; see Mohr’s label notation in 
Fig. 1). Smith (881b) recalled, “I think | must be the discoverer as I know of no 
one else who has ever collected any of the material... It makes almost 
inpenetrable “Privet Thickets” as they are called, and the peculiar aroma of the 
leaves as you feel your way through, is exceedingly characteristic. | have not 
seen the plant anywhere else, than along the banks of the Cahaba River where 
it grows upon limestone much like Cedar.” 


From press to print 
In January 1878, Smith approached Charles Theodore Mohr (1824-1901) about 
preparing a mounted, organized herbarium for the Geological Survey from 
Smith’s collections. Mohr’s poor health and business obligations put the project 
on hold until May 1879, when he received the plants (Mohr 1879a). By July the 
Croton had come to Mohr’s attention and he directed an inquiry to Smith: 
“Amongst some other interesting plants collected at Pratts Ferry (vi. Forrestonia [sic], Forrestiera 
ligastrina [sic] et al.) | find a shrub marked “Privet.” Is it a truly indigenous plant or escaped from 
cultivation[?] It is very near Eleagnus Canadensis (the specimens being without flowers or fruit | 
cannot decide positively). It should be very a indeed to find that Northern shrub at once com- 
ing up afar in Central Ala{[bama].” (Mohr 187¢ 
The specimen (Fig. |) described by Mohr in this letter and annotated by him 
“Type Spm. [specimen] is apparently Smith’s earliest collection previously noted; 
it is in young bud. Mohr later recalled, “I found first a sprigg [sic] of it amongst 
the plants of the Geol. Survey of Ala. collected by the Professor [Smith] not- 
withstanding the poverty of the specimens bearing only a male spike with the 
flowers yet in the bud. I recognized as plant entirely new” (Mohr 1882a). 

After examination of a single male flower, Mohr tentative oe referred the 
Croton to section Hendecandra Eschw. and sent the specimens off to George 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 475 


Engelmann (1809-1884) with a cover letter dated February 24, 1880, noting 
that “perhaps the plant is known to you from Texas or N. Mexico” (Mohr 1880b). 
Mohr wrote to Smith, “I was somewhat astonished to find the Croton (Privet) by 
Dr. Engelmann referred to Cr. argyranthemum [Croton argyranthemus Michx,], 
a straggly herb not infrequent in our dry pine barrens here... It will be neces- 
sary to secure large specimens with a portion of the lower stem to establish its 
character as a true shrub and in full bloom with male and female flowers, the 
plants appear to be dioecious” (Mohr 1880c). Little progress had been made by 
October 11, 1880, when Mohr reported to Smith, “the Croton continues to be [a] 
riddle to all who get hold of it” and lamented the lack of adequate fertile mate- 
rial. The lack of fertile material explains his belief that the plant was dioecious 
(it isactually moneocious, with typical bisexual Croton inflorescences), though 
by this time he had the remnants of female flowers (Mohr 1880d). Mohr recalled, 
“Some time later I was so fortunate to find a unripe coped and a female flower, 
from which it was evident that it has its pl [sic] the section Eleuteria” 
(Mohr 1882a). There is no mention of the Croton in the Alabama floristic check- 

list Mohr and Smith published in 1880 (Mohr 1880a). 

Mohr turned to Alvan Wentworth Chapman (1809-1899), who was then 

involved in the preparation of a second edition of his Flora of the Southern 
United States (ie. sau 1883a). Mohr (1881la) wrote to Smith: 


] ] 


for the * roton.Can you not geta slip for me to plantin my 
gar iene I have sent at his request all of the material I had received of yours (except that what I oad sent 


before to Prof. Engelmann who certainly took a wrong view of it) to Dr Chapman. He was anxious to 
describe the plant in the new edition of his Flora, he agreeing with me at first sight that [it] is see 

In the pursuit of more material, Chapman wrote directly to Smith, who replied 
April 8, 1881: 


“The croton you mention is a shrub he height of 8 to 10 feet. [have seen some 
ae three puenes in amiclee ie nae an aa ee thicket, and the aromatic odor 

i ility [probably Jacob S. Hansberger, 
see below] who has nronsed to get specimens du po oa of the year so that | may be 


able to get sufficient material for eee Dr. Mohr WG now all my material. When | get other 
specimens, I will remember to send you some” (Smith 1881la). 

Chapman requested all the material Mohr had “on hand,” and Mohr reported to 
Smith that the request had been honored including a specimen with “ripened 
fruit in perfect state” (possibly this collection is US-956957, see below) (Mohr 
188la, 1881b). In a letter dated July 10, Smith (1881b) reported that he was ex- 
pecting new material in a “week or two,” presumably from his “correspondent.” 
He also approved Chapman's proposed name “Croton alabamense” (Lc.). On July 
17, Smith was able to send Chapman “a small box containing specimens of Cro- 
ton alabamense, in different stages of its growth.” He continued, “From these, I 
hope you will be able to get all the information heretofore lacking. You might 
plant some of the seeds and perhaps raise it. | have several flourishing plants in 


476 BRIT.ORG/SIDA 22(1) 


flower pots, which I have lately received from Pratt’s Ferry” (Smith 1881c). These 
additional specimens proved sufficient for the complete description. The “18 
inch seedlings” received from the “correspondent” in July 1881, Smith planted 
in his yard and they attained a height of “7 feet” by 1889 when Mohr reported 
on their development (Mohr | 

Chapman traveled north, ne with Charles Mohr while passing though 
Washington (Mohr 1882b), and delivered the manuscript for his Flora to the 
printer in the last week of July 1882 (Chapman in Oakes 1932). Proof correc- 
tions continuing into the fall and the preface was dated December 26, 1882 
(Chapman 1883a). Copies of the Flora were circulating by February 1883 (Chap- 
man 1883b). The main body of the Flora was merely a corrected version of the 
first edition (Chapman 1860) and new taxa, including the Croton, were con- 
tained in an appended supplement. Chapman (Lc.) credited Smith as the au- 
thority for the species but added “ined.,” an amendment finally removed in the 
entirely revised third addition of the Flora (Chapman 1897). 

Mohr was preoccupied with work for Charles Sprague Sargent (1841-1927) 
on forest trees for the Tenth Census of the United States and could not visit Bibb 
County until November 1882 (vouchers at A,GH, MO and US variously give the 
date as November 11 or 12). Mohr located the Croton while collecting wood samples 
of Quercus durandii Buckley on his final trip for Sargent. Of the trip Mohr notes: 


“About the middle of last month I was over on the little Cahaba river near Tionus [sic]. 1 went by way 
of Montevallo; having to return the same day I made 32 miles on horseback, arriving at my quarters 


Mr. Aldriches at 10 o'clock pm... At the time of my visit to the ‘Privet’ thickets near Tionus the plant 
was just putting forth the flower buds of the staminate spiklellets, which as Mr. Hansberger informs 
me open with the first warm days of le spring. Nota ve a of evenarudimentary pistillate [lower 


could be found and the fruit of this season was entirely gone... Mr. Ha tl received me with the 
genuine warm hospitality of the es n gentlemen, went oe me through the woods and gave 
me all assistance in his power.. 1am bound to visit again this locality so aes on [sic] botanical trea- 
sures” (Mohr 1882c). 

Tionis was a community located along lower Six Mile Creek (sec. 26, T24N, RLOE; 
fide Smith 1870-1889) and readily evident on period maps (see online histori- 
cal map archive at the University of Alabama, http://alabamamaps.ua.edu/ 
index.html). A post office was established in 1870 and Jacob S. Hansberger (d. 
1887) served as postmaster from 1874 until 1887 when it was discontinued prior 
to his death (Stewart 1983). Bibb County deed books show that he owned over 
200 ha of land in the vicinity (accrued as many parcels over a period of years). 
Smith knew Hansberger from his Bibb County trips and they had explored Six 
Mile Creek together shortly after Smith first discovered the Croton (fide entry 
for August 1, 1877; Smith 1870-1889). It is probable that Smith instilled in 
Hansberger an interest in the Croton and that he served as Smith’s “correspon- 
dent” and local expert. The exact locations of Tionis and Hansberger’s house 
are presently uncertain and | have not been able to assess the patchy Croton 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 477 


distribution in relation to them. Even today it is clear that Croton is readily avail- 
able in the general area without having to travel all the way to Pratt’s Ferry (ca. 
6.4 km to the west). 

In the fall of 1882 Frederick Hoffmann, editor of Pharmaceutische 
Rundschau, requested Mohr to submit an article on Croton alabamensis for that 
journal (Mohr 1882b). Mohr had noted that the Croton bark was of interest to 
“pharmacognosy” as a possible native substitute for cascarilla that came from 
the bark of the West Indian Croton eluteria (L.) Sw. Mohr decided to defer writ- 
ing the article until he could obtain flowering material to clear up the “point of 
the sexual relation of their flowers” and have a photograph of the plant taken. 
In March 1883 Smith sent Asa Gray a print of the photograph (annotation on 
back of print says the photo was taken February 28, 1883; Smith 1883) and Mohr 
received a copy in May (Mohr 1883a). In June 1883, while collecting wood for 
the Louisville Cotton Exposition, Mohr returned to Pratt’s Ferry, ostensibly for 
Quercus durandii wood, but also unsuccessfully for Croton seeds and flower- 
ing specimens. (Mohr 1883p). 

Mohr’s labors finally bore fruit. In a paper read in January 1884 before the 
Mississippi Valley Horticultural Society, he presented his views on the possible 
use of Rhus cotinoides (= Cotinus obovatus Raf.), Neviusia alabamensis A. Gray, 
and Croton alabamensis as horticultural subjects (Mohr 1884). The same three 
species were treated in an expanded form by Mohr for the January 1887 issue of 
Pharmaceutische Rundschau (Mohr 1887). An etching of a cultivated seedling 
was substituted for the originally intended photograph. Preparation for this 
article included the examination of additional blooming material that was col- 
lected by Smith in his garden and received by Mohr in April 1886 (Mohr 1886). 
This material probably included a collection in the Mohr Herbarium dated April 
16,1886 from “my yard (Tuscaloosa), transplanted from Pratt’s Ferry” (E.A. Smith 
s.n., UNA-00020895, annotated by Mohr “Type Spm.”). In 1888, Mohr’s prepara- 
tion of an article for Garden and Forest brought renewed questions to Smith on 
the life history of the Croton and requests for specimens (Mohr 1888). Mohr 
hoped to have the article sent off to press by the end of August 1888 and it ap- 
peared in the December 11, 1889 issue of Garden and Forest. In March 1889 Smith 
sent seedlings of the Croton to Sargent and Mohr (Smith 1889). 


Typification and historical collections 

Chapman gave no collection data beyond “central Alabama’ in his original de- 

scription, therefore a lectotype is selected here: 

Croton alabamensis E.A. Sm. ex Chapm., Flora of the Southern United States, 
ed. 2:648. 1883. Lectotype: UNITED STATES. ALABAMA: E.A. Smith s.n.(US-935923, ex John 
Donnell Smith herbarium). Other original material: Alabama, 4 Apr 1881 (date given on 
supplemental tag attached to stem), E.A. Smith s.n. (MO-1904803); Alabama, E.A. Smith s.n. 
(US-956957, ex Biltmore Herbarium). Alabama, Tuscaloosa, E.A. Smith s.n. (F-99491, ex H.N. 
Patterson herbarium; MO-784596, ex Chapman estate from 1899 purchase by MO). 


478 BRIT.ORG/SIDA 22(1) 


Herbaria (GH, MO, NY, UNA, and US) containing the best representation of 
collections from Chapman, Mohr, and Smith were searched for historical Cro- 
ton collections. Five Croton specimens were located that bear evidence of hav- 
ing been seen by Chapman (ie., bear writing in Chapman’s hand), although 
additional specimens may turn up in his widely scattered exciccatae (see Stafleu 
and Mennega [1997] for a listing of the 27 herbaria with Chapman material). It 
is unclear whether the two sheets labeled “Tuscaloosa” by Chapman should be 
considered original material. If taken asa literal exact locality then the collec- 
tions came from cultivated material and most likely post-date the preparation 
of the original description (i.e., post 1881). Smith did cultivate plants in Tusca- 
loosa that were eventually a source of specimens, but these were probably the 
small, valuable seedlings also received in the summer of 1881, and not likely to 
become herbarium fodder at that time. On the other hand, label locality data 
secondarily penned by Chapman and not the collector might be distorted or 
incomplete. Chapman did generalize that Pratt’s Ferry was “near Tuscaloosa” 
(see below) or he may have referred to Smith being based in Tuscaloosa. 

John Donnell Smith (1829-1928) corresponded with and received speci- 
mens from Chapman during at least 1884-1886, based on specimen provenance 
notations in Donnell Smith’s hand and fragments of Chapman letters attached 
to herbarium specimens at US. Donnell Smith’s personal herbarium (including 
the lectotype, see Figure 2 with embossed stamp) and library were donated to 
US in 1905. The lectotype has an attached undated fragment of a Chapman let- 
ter (consistent in the distinctive paper with other letter fragments established 
as dating from 1884-1886), presumably to Donnell Smith, that reads: 


“Croton Alabamensis is not found along R{ailJRoads and being a shrub will remain pretty quiet in its 
secluded home near Tuskaloosa [sic]. Strange that it should be in flower every month from Sept to 
May through the coldest weather and yet they sent me specimens collected in all these months!” 
Based on the detailed historical account given above, it appears that Chapman 
had material from two collectors (E.A. Smith and J. Hansberger), multiple dates 
and perhaps several Bibb County localities on hand for his original diagnosis. 
The circumstantial evidence (ie., Smith did not visit the area between the time 
of the Croton discovery and publication, and he also refers to receiving speci- 
mens from a “correspondent”) suggests that much of the material was collected 
by Jacob Hansberger near Tionis, and the lectotype may have been collected by 
him in April 1881 and sent to Smith, who forwarded the specimens to Chap- 
man for description. The two branches on the lectotype sheet have young fruits 
at slightly different developmental stages and may come from separate late 
spring collections. Of the five sheets of original material, only the selected lec- 
totype contains an additional printed label that indicates the specimen was 
associated with the “Flora of the Southern United States and Supplement’ (ie, 
Chapman 1883a). 

In addition to sending duplicates to correspondents, Chapman sold sev- 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 


U.S. National Herbequm. det 


00730774 pea Mee, 


TOTYPE: 
Croton alabamensis E. A. Smith ex Chapman 
Fl. S. U.S. ed. 2: 648 


Ky Wolk 17-2.0904 


TT ne 


, 1883 


Cee rea Pig ins, EX HERS. / United Stat 
" and Supplement, A. W. Chapman, M.D.” 
: sae how ar Sane -- Brett: 
Af 
‘ nee ~ tte 
bs af 
V2 yore Pad yb ry rieee 


JA 


Loulin Mabivrrrecrdit, F Kidugy 


Bias ate 
VE Dop Mir Slat ri 


aeate 


alqhamensic(F A S, 


Fic. 2.Crot I 


“ 


Chapman. See text for transcription of letter. 


480 BRIT.ORG/SIDA 22(1) 


eral sets of specimens that are massive enough to have been described as “her- 
baria.” However, he always retained a personal working collection, the residue 
of which was purchased from his estate by MO in 1899. In 1896 Chapman se- 
lected and sold the best set of material on which his 1883 Flora was based, to 
the Biltmore Herbarium in Asheville, North Carolina. The Biltmore Herbarium, 
modeled after the Royal Botanic Gardens, Kew, was established ca. 1894 for 
George Vanderbilt (1862-1914) asa scientific adjunct to his lavish Biltmore Es- 
tate. At the time of the Chapman purchase, the herbarium was trying to estab- 
lish itself as one of the foremost botanical institutions in the southern United 
States through purchasing and exchanging collections, hiring of collectors, and 
publishing the eponymous scientific journal Biltmore Botanical Studies. The 
enterprise withered after about 10 years of activity (peaking ca. 1901-1903) due 
to Vanderbilt’s financial downturn and untimely death. A 1916 flood destroyed 
three-quarters of the collection and the residue of ca. 25,000 sheets was given 
to US in 1917. The Chapman collection was reportedly stored in a “vault” and 
largely salvaged (Maxon 1917; Boynton 1936). Only one flood-damaged fruit- 
ing specimen (US-956957, stamped ex Biltmore Herbarium) of Croton 
alabamensis clearly has this provenance and was used at the Biltmore in an 
illustration (see Lounsberry 1901:306). It is possible that other C. alabamensis 
specimens, including other type material at Biltmore, were destroyed. 

An anomalous Tennessee record (Tullahoma, Coffee Co., 10 Aug 1899, TG. 
Harbison 725, NCU-9391) has connections with Biltmore. Thomas Grant Harbison 
(1862-1936) was a member of the Biltmore Herbarium staff and made extensive 
collections of the southern flora. His incomplete Biltmore field notes deposited 
at US (partial number series of duplicate field notes bracketing, but not includ- 
ing, the Croton collection in question) place him in Tullahoma, but suggest errors 
on the specimen label which was made long afterwards (ie., Harbison 725 would 
have been collected on 12 Aug 1899 and not 10 Aug as reported with the speci- 
men). A Croton alabamensis collection with the same collection number and from 
a well-established locality “On sandstone bluffs along the Warrior River, Tusca- 
loosa, Alabama,” T. G. Harbison 725, 11 Oct 1911, A) may have been a source of 
confusion. The credibility of the Tennessee record is doubtful. 


ACKNOWLEDGMENTS 


I thank the curators and staff of the noted herbaria and archives for access to 
their collections and permission to quote archival materials. | am grateful to 
LJ. Davenport (Samford University) for the Smith to Chapman typescripts and 
to B. van Ee for unpublished information on his recent genetic studies. J. Alli- 
son, LJ. Davenport, LJ. Dorr (US), and J. Reveal (emeritus MARY) provided help- 
ful comments on the manuscript. At the Geological Survey of Alabama, A. 
Sartwell provided information on Smith and alerted me to the archives formerly 
in his care and LS. Dean clarified Smith’s geological terminology. 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 481 


REFERENCES 


A.uison, J.R.and T.E. Stevens. 2001.Vascular flora of Ketona Dolomite outcrops in Bibb County, 
Alabama. Castanea 66:154-205 

Aptet, G.H.,R.D. Laven, M.B. FALKNer, and R.B. SHaw. 1994. Population and site characteristics of 
a recently discovered disjunct population of Croton alabamensis (Euphorbiaceee). Sida 
16:37-55. 

Berry, PE., A.L. Hipp, K.J. Wuroack, B. Van Ee, and R. Rina. 2005. Molecular phylogenetics of the 
giant genus Croton and tribe Crotoneae (Euphorbiaceae sensu stricto) using ITS and 
trnL-trnF DNA sequence data. Amer. J. Bot. 92:1520-1534. 

Boynton, F.E. 1936. Letter to C. D. Beadle, 28 October, from Old Fort. Library of the Gray 
Herbarium, Harvard University. 

CHapman, A.W. 1860. Flora of the Southern United States. Ivison, Phinney & Co., New York. 

CHapman, A.W. 1882. Letter to A. Gattinger, 3 August.|In: Oakes, H.N. 1932.A brief sketch of 
the life and works of Augustin Gattinger. Cullom & Ghertner Co. Nashville, Tennessee. 

CHAPMAN, A.W. 1883a. Flora of the Southern United States. 24 ed. Ivison, Blakeman, Taylor, 
& Co., New York. 

CHapman, A.W. 1883b. Letter to G. Engelmann, 6 February, from Apalachicola. Missouri Bo- 
tanical Garden Archives. 

Cuapman, A.W. 1897. Flora of the southern United States. 3'¢ ed. Cambridge Botanical Sup- 
ply Co., Cambridge, Massachusetts 

Davenport, L.J. 1979. Charles Mohr and plant life of Alabama. Sida 8:1-13. 

Davenport, L.J. 1988. Charles Mohr, botanist. Alabama Heritage No. 10:32-45. 

ELuison, R.C. 1984, Bibb County Alabama. The first hundred years, 1818-1918. The Univer- 
sity of Alabama Press. 

ELuison, R.C. 1993. Place names of Bibb County: Abercrombie to Zuzu. Cahaba Trace 
Commision, Brierfield, Alabama. 

Farmer, J.A. 1962. An ecological life history of Croton alabamensis E.A. Smith ex Chapm. 
Unpublished Ph.D. dissertation, University of Alabama, Tuscaloosa. 

Farmer, J.and J.L.THomas. 1969.Disjunction and endemism in Croton alabamensis.Rhodora 
71:94-103. 

GEOLOGICAL SURVEY OF ALABAMA. 1894. Geological map of Aleaiia Bien & Co., New York. 
Avaialbale at http://alabamamaps.ua.edu/historicalma| gy/index.html. 

GinzBarG, S. 1991. Croton alabamensis Chapm. (Euphorbiaceae) ane: populations in 
Texas. Unpublished M.A. thesis, University of Texas, Austin. 

GinzparG, S. 1992. A new disjunct variety of Croton alabamensis (Euphorbiaceae) from Texas. 
Sida 15:41-52. 

Harper, R.M. 1906a. Diary entry of 15 February. Harper Papers. W.S. Hoole Special Collec- 
tions Library, University of Alabama. 

Harper, R.M. 1906b. A December ramble in Tuscaloosa County, Alabama. Pl. World 9:102, 
104-107. 


482 BRIT.ORG/SIDA 22(1) 


Lounsserry, A. 1901.Southern wild flowers and trees. Frederick A.Stokes Company, New York. 

Maxon, W.R. 1917.Memorandum to R. Rathbun, 5 June, from Washington. Registrar's files 
for Biltmore Herbarium accession 61193, Smithsonian Institution. 

Monk, C.T. 1879a. Letter to E.A. Smith, 21 May, from Mobile. W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.T. 1879b. Letter to E.A. Smith, 29 July, from Mobile.W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.T. 1880a. Preliminary list of the plants growing without cultivation in Alabama, 
from the collections made by Eugene A. Smith, Tuscaloosa, and Chas. Mohr, Mobile, 
Ala. Compiled by Charles Mohr. No publisher. 

Mone, C.T. 1880b. Letter to G. Engelmann, 24 February, from Mobile. Missouri Botanical 
Garden Archives. 

Mor, C.T. 1880c. Letter to E.A. Smith, 22 March, from Mobile. W.S. Hoole Special Collec- 
tions Library, University of Alabama. 

Mone, C.T. 1880d. Letter to E.A. Smith, 11 October, from Mobile. W.S. Hoole Special Collec- 
tions Library, University of Alabama. 

Mone, C.T. 1881 a. Letter to E.A. Smith, 27 May, from Mobile. W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.T. 1881b. Letter to E.A. Smith, 15 July, from Mobile. W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.1. 1882a. Letter to A. Gray, 5 June, from Mobile. Library of the Gray Herbarium, 
Harvard University. 

Mone, C.T. 1882b. Letter to EA. Smith, 5 July, from Washington. W.S. Hoole Special Collec- 
tions Library, University of Alabama. 

Mone, C.T. 1882c. Letter to E.A. Smith, 15 December, fromm Mobile. W.S. Hoole Special Col- 
lections Library, University of Alabama. 

Mone, C.T. 1883a. Letter to E.A. Smith, 22 May, from Mobile. W.S. Hoole Special Collections 
Library, University of Alabama. 

Monr, C.T. 1883b. Letter to E.A. Smith, 19 June, from Mobile.W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.T. 1884. Rare and little known trees and shrubs of Alabama. Trans. Mississippi 
Valley Hort. Soc. 2:216-29, 

Mone, C.T. 1886. Letter to E.A. Smith, 22 April, from Mobile. W.S. Hoole Special Collections 
Library, University of Alabama. 

Mone, C.T. 1887.Ueber drei vereinzelte Burger des Floren-Gebietes der nordamerikanischen 
Sudstaaten, Pharm. Rundschau (Berlin and New York) 5:8—11. 

Mone, C.T. 1888. Letter to E.A. Smith, 25 August, from Mobile. W.S. Hoole Special Collec- 
tions Library, University of Alabama. 

Mone, C.T. 1889. The la[te]st addition to the shrubs of eastern North America. Gard. & 
Forest 2:592, 594. 

Mour, C.T. 1898. Letter to A.M. Ferguson, 19 April, from Mobile. Missouri Botanical Garden 
Herbarium (attached to specimen of Croton alabamensis). 


WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 483 


Mour, C.T. 1901. Plant life of Alabama: an account of the distribution, modes of associa- 
tion,and adaptations of the flora of Alabama, together with a systematic catalogue of 
the plants growing in the state. Contr. U.S. Natl. Herb, 6:1-921. 

Smith, E.A. 1870-1889. [WPA? typescripts of 1936]. Field notes of E.A.Smith.v. 1:1870-1875, 
Vv. 2:1876-1882, v. 3:1883-1889. Library, Geological Survey of Alabama. 

Smith, E.A. 1881a.Letter to A.W. Chapman, 8 April, University of Alabama. [WPA? typescript] 
Library, Geological Survey of Alabama. 

SmitH, E.A. 1881b. Letter to A.W. Chapman, 10 July. [WPA? typescript] Library, Geological 
Survey of Alabama. 

Smith, E.A. 1881¢. Letter to AW. Chapman, 17 July. [WPA? typescript] Library, Geological 
Survey of Alabama. 

Smitu, E.A. 1881d.Letter to A.W. Chapman, 7 September, from Prattville. [WPA? typescript] 
Library, Geological Survey of Alabama. 

Smito, E.A. 1883. Letter and photograph to A. Gray, 11 March, from Tuscaloosa. Gray Her- 
barium (filed with Croton alabamensis), Harvard University. 

Smith, E.A. 1889. Letter to C.S. Sargent, 26 March, from University, Alabama.Gray Herbarium 
(filed with Croton alabamensis), Harvard University. 

Smith, E.A. 1907. Letter to R.M. Harper, 8 September, from University, Alabama. Harper Pa- 
pers. W.S. Hoole Special Collections Library, University of Alabama. 

Starteu, FA. and E.A. MenNeca. 1997. Taxonomic literature. Supplement IV: Ce-Cz. Koeltz 
Scientific Books, Germany. 

Stewart, F.R. 1983. Alabama's Bibb County. Unpublished Doctor of Education dissertation, 
University of Alabama. 

vAN Ee, B. and PE. Berry. 2005. The Caribbean-centered Moacroton clade within Croton 
(Euphorbiaceae sensu stricto).Abstract. Available at:http://www.2005.botanyconference.org 

VAN Ee, B., N. Jevinski, PE. Berry, and A.L.Hiee.In press. Phylogeny and biogeography of Croton 

abamensis (Euphorbiaceae), a rare shrub from Texas and Alabama, using DNA se- 
quence and AFLP data. Molec. Ecol 

Weester, G.L. 1967.The genera of Euphorbiaceae in the southeastern United States. J. Ar- 
nold Arbor. 48:303-430. 

Weester, G.L.M.J. pet Arco Acuitar, and B.A. SmitH.1996. Systematic distribution of foliar tri- 
chome types in Croton (Euphorbiaceae). Bot. J. Linn. Soc. 121:41-57 

Weesster, G.L. 1993. A provisional synopsis of the sections of the genus Croton 
(Euphorbiaceae). Taxon 42:793-823. 

Worrorp, B.E. and E.W. CHester. 2002. Guide to the trees, shrubs, and woody vines of Ten- 
nessee. The University of Tennessee Press. 

Wotre, S.R. 1983. The University of Alabama. A Pictorial History. The University of Alabama 
Press. 

Wuroack, K.J., P. HorFMANN, and M.W. ee 2005. Molecular phylogenetic analysis of 
uniovulate Euphorbiaceae (Euph ceae sensu stricto) using plastid rbcLl and trnL-F 
DNA sequences. Amer. J. Bot. 92:1397-1420. 


=e 


484 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Mark Turner and Puytis Gustarson. 2006. Wildflowers of the Pacific Northwest. 
(ISBN 0-88192-745-7; 978-0-88192-745-0, flexibind). Timber Press Inc, 133 
S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
www.tim berpress.com, mail@timberpress.com, 503-227-2878, |-800-327- 
5680, 503-227-3070 fax.). $27.95, 512 pp., 1248 color photos, 1221 maps, 31 
line drawings, 5 3/4" x 81/4". 

More than 1200 native and non-native ee are described and illustrated, including 1247 color 

photos and 1219 ran ( distribution). The region covered is all of Washing- 

ton and Oregon, a: Seurhiwectern Br itish Columbia and the northern third of California. An 

ecoregions map and excellent ey intro een nae “C sas Geography, and Plant Habitats,” set 


the stage. The photos lowers as well as critical vegetative fea- 


tures, and the morphological and meee notes are ae and to the point. Six categories of flower 
color (white, yellow, orange, pink to red or red-purple, violet to oe or blue- purple, brown and green) 
provide the first division in the arrangement of species, followed wit} ‘h color category by ma- 


jor structural features (e.g, number of petals, petals forming a a or not, ovary inferior or superio 
composite flowers). My perennial response to “color-coded” taxonomy is that so many people ene 
this book will wish the blue- ene Ww Bite: flowered (etc.) species of a single genus were together, but 
apart from this personal bias, this ! “third-generation” field guide, about as good as they 
get, useful to everyone interested in the flora of this region.—Guy Nesom, Botanical Research Institute 
of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


SIDA 22(1): 484. 2006 


TWO NEW SPECIES OF ELYMUS (POACEAE) IN THE 
SOUTHERN U.S.A. AND OTHER NOTES ON 
NORTH AMERICAN ELYMUS SPECIES 


Julian J.N. Campbell 


The Nature Conservanc cy ( (} entucky Chapter) 
42 leas ae Street 
Lexington, b 40508-2018, U.S.A. 


ieamipbel@tne org 


ABSTRACT 


wo new species of Elymus L. are described, E. churchii JJ.N. Campb. and E. texensis J.J.N. Campb. 
Elymus churchii grows in the Ouachita and Ozark Mountains of Arkansas, Missouri and Oklahoma. 


It is similar to E. svensonii G.L. Church, but differs in its lemmas, which are usually pubescent, and 
onger awns; its spikelets, which have fewer florets; its ienicno des, which are longer, wit 
teed dorsal ridges and green longitudinal bands; and its foliage, which is typically less pubescent 
and less glaucous. Elymus texensis grows on the Edwards Aue in southwest Texas. It is similar to 
E. pringlei Scribn. & Merr, but differs in its larger anthers; its glabrous lemmas; its larger spikelets; 
its longer spikes, with ee internodes that are esas with oe longitudinal bands, and gener- 


ally glabrous; and its foliage, which is ly pilo e. The new succes are sais from 


yi ] 
£ 


few collections, and need much more cee to assess their 
bution. They are published here to allow inclusion in the Flora of North America, pone 24, and to 
stimulate further attention to them. Some additional notes and corrections are appended regarding 
the author's previous publications on Elymus. 


RESUMEN 
Se describen dos nuevas especies de Elymus L., E. churchii J.J.N. Campb. y E. texensis ne Campb. 
Elymus churchii vive en las montafias Ouachita y Ozark de Arkansas, Missouri y Oklahoma. Es simi- 
lt a E. svensonii — L. Church, p ncia por sus es que son penal pubes ntes, y 
ill u is, que son 


mas poe con eostillac derales Rigpidas y bandas verdes longitudinales; y su - llaje, que es 
tipicamente menos pubescente y menos glauco. Elymus ene vive en el Edw a Plateau en el 
sureste de Texas. Es saaues a E. ae 


oO 
] ] . ll a a 


aq n 


as fe con Bande verdes longitudinale y pene erie glabras; y su follaje, ae es a veces 
ee piloso. Las nuevas especies se conocen de pocas colecciones, y necesitan mucho mas 
estudio para evaluar su estatus gent y distribucion geografica. Se publican aqui para que 


nee incluirse en mila Flora de Norte América, volumen 24, y para estimular la atencion sobre ellas. 
Se anaden les y correcciones a Sree previas del autor sobre Elymus 
INTRODUCTION 


In 2002, I provided descriptions, specimen data, and distribution maps for two 
taxa of Elymus that seemed different from other members of the genus with 
paired spikelets and reduced or vestigial glumes (Campbell 2002a). I refrained 


SIDA 22(1): 485 — 494. 2006 


486 BRIT.ORG/SIDA 22(1) 


then from formal taxonomic recognition, but after further revison | am now 
more confident. To allow inclusion in the Flora of North America, volume 24 
(M.E. Barkworth et al. in prep.), | present below the two new names, anda key to 
the species of Elymusthat have paired spikelets and reduced or vestigial glumes. 
There has been much uncertainty in the circumscription and phylogeny of 
these species, which would have been grouped together under the broad con- 
cept of Elymus interruptus Buckley by Hitchcock and Chase (1951). 


Elymus churchii J.).N. Camp., sp. nov. (Fig. 1). Type: US.A. ARKANSAS: Conway Co: Petit 
Jean State Park, rocky bluffs, PO. Morrilton, 1500 ft, 3 Jul 1957, D. Demaree 37234 (HOLOTYPE: 
UARK:; IsoTyPES: OKL, SMU-BRIT). 


Affine Elymo svensonii G.L. Church sed differt lemmibus plerumaque pubescentibus aristis (10-)20- 


30(-35) mm longis, spiculis flosculis 3(-5), internodis spicarum (5-)7-13(-18) mm longis viridi-vittatis 


dorsaliter hispidis, foliis paginis adaxialibus glabris vel pilosis, et plantis non glaucissimis. 

Plants cespitose, of ten somewhat, but not strongly, glaucous. Culms 50-120 cm, 
erect; nodes usually 4-8, exposed or covered, often reddish-brown or blackish, 
glabrous. Leaves evenly distributed; sheaths usually glabrous, or sometimes 
pubescent at the summit; auricles 1-2 mm, often reddish brown or blackish; 
ligules up to 1 mm, often reddish-brown; blades 3-11 mm wide, lax, adaxial sur- 
faces glabrous or short-pilose. Spikes 10-18 cm long, 3-5 cm wide, slightly nod- 
ding, with 2 spikelets per node; internodes (5-)7-13(-18) mm, about 0.2 mm thick 
at the thinnest sections, flexuous, with green longitudinal bands along the con- 
cave sides, glabrous except for the two hispid dorsal ridges; disarticulation be- 
low each floret, but not below the glumes. Spikelets usually appressed, 10-15 
mm (excluding awns), with 3(-5) florets. Glumes often unequal (differing in 
length by more than 5 mm), or sometimes vestigial to absent from the upper 
spikelets or throughout, 0-15(-20) mm long including the undifferentiated 
awns, indurate at base, 0.1-0.3 mm wide, setaceous to subulate, with 0-1 dis- 
tinct vein, glabrous, awns often outcurving; lemmas 8-10 mm, pubescent, or 
occasionally glabrous, awns (10-)20-30(-35) mm, slightly to (at maturity) 
strongly outcurving; paleas 7-9 mm, obtuse to truncate, or emarginate; anthers 
about 2.5-3 mm, evident in June. 2n = unknown. 

Elymus churchii grows in dry rocky soils, often base-rich, in open woods 
on ridges, bluffs and river banks of the central Ouachita Mountains and the 
western Ozark Mountains, in Arkansas, Oklahoma and Missouri. It was previ- 
ously included within a broad concept of E. interruptus (Steyermark 1963; Smith 
1991). It is similar to E. svensonii, which is disjunct in Kentucky and Tennessee. 
It might reasonably be combined as a subspecies or variety of F. svensonii, but 
differs in its lemmas, which are usually pubescent, and have longer awns; its 
spikelets, which have fewer florets; its rachis internodes, which are longer, with 
hispid dorsal ridges and green longitudinal bands; and its foliage, which is typi- 
cally less pubescent and less glaucous. It may have originated, like E. svensonii, 


CAMPBELL, 


tii | 


MG ck. Sah calek he 
D 


ea 
ZN 


\ eee 


~~, 
os 


eo ~~ 
aw, 


ith 
v7 % 


a 
———_ 


=. 
Fao even Stl Op 


haa 


a 
oo 


Fic. 1 Ely hurchii (d f Demaree 29337 unless noted). A. Habit. B. Upper | i f cul itt pil 
viewed on plane with alternating spread of spikelets (B2 from Palmer 57957). C. Sheath summit and blade bases. D. 
hdaviallesfeurtace- showing Weer ey . ‘al (showing variation in size), viewed 
in pl pikelet spread (with abaxial view of Igl pikelet, and largely side view of lateral glume). F. 
Spikelet, with lateral view of florets (F2 from Palmer 57957). G. Mature floret in abaxial view (| ft) and adaxial vi 
(right). H. Cross-secti f mature, indurate gl I (showi 
internode. 


J 


ize). I. Cross-section of central rachis 


488 BRIT.ORG/SIDA 22(1) 


from introgression of E. canadensis L. and E. hystrix L., and there are occasional 
specimens suggesting transitions to both species (Campbell 2002a). This spe- 
cies is named after George L. Church, who conducted several useful studies of 
Elymus in eastern North America, focusing on problems of hybridization and 
the species with reduced glumes (Church 1967). 

ParatyPes: ARKANSAS. Baxter Co.: rocky shaded blulf, White Rv. PO. Lakeview, Bull Shoals Dam 
Reservoir, 600 ft, 1 Jul 1950, D. Demaree 29337 (OKL, TEX). Logan Co.: Magazine Mt, about 45 mi east 
of Fort Smith, 8 Aug 1942, D.M. Moore 420118 (TEX), Sep 1947, D.M. Moore 470639/642 (UARK, NCU, 
US), and 30/31 Jul 1949, D.M. Moore 490422/441 (UARK). Newton Co.: Big Bluff above Buffalo Rv, 5 
mi below pes: 5 Jun 1953, D.M. Moore 53259 (UARK). OKLAHOMA. Le Flore Co.: Rich Mt., roadside. 
oak-] nant, 16 Jun 1940, H. Taylor 110 (OKL); see also See ener 


ihe salves collection known from Missouri has relatively short rachis intenodes 
(ca. 5-7 mm): Christian Co., 3 miles west of Nixa, rocky wooded banks, 24 Jun 
1954, E.J. Palmer 57957 (SMU). I previously suspected that all of Steyermark’s 
(1963) records of Elymus diversiglumis Scribn. & CR. Ball or E. ee aa in 
Missouri could be grouped with the plants described here as E. churchii(Camp- 
bell 2002a). However, some of these records were based on misidentifications: 
EJ. Palmer 66416 (UMO) is E. glabriflorus var. australis (Scribn.&@ CR. Ball) JJ.N. 
Campb.; EJ. Palmer 55939 (UMO) is E. hystrix var. bigelovianus (Fernald) 
Bowden. 


Elymus texensis J.J.N. Camp, sp. nov. (Figs. 2a, b). Type: US.A. TEXAS. Gillespie Co: Ser- 
pentine Mounds, about 9 mi N of Willow City, hilly area vegetated mainly with grasses, 18 
May 1906, E.S. Nixon 531 (HOLOTYPE: TEX 5322) 


Affine Elymo pringlei Scribn. & Merr. sed differt antheris 4.5-6 mm longis, lemmil s glabris, spiculis 
25-40 mm longis flosculis 5-8, 3 icis 9-20 cm longis phauaue (5- ce 15(- ens mim longis an 


vittatis glabris praeter margi _foliis pag hirsutis vel dense 
brevipilosis. 

Plants cespitose, glaucous. Culms 70-110 cm, erect; nedes usually 4-6, mostly 
exposed, glabrous. Leaves evenly distributed; sheaths glabrous; auricles about 
0-1 mm, sometimes adherent to sheath summit, pale to purplish brown; ligules 
about | mm, erose; blades 2-9 mm wide, lax, or somewhat involute, adaxial sur- 
faces thinly scabrous-hirsute or densely short-pilose. Spikes 9-20 cm long, 2- 
25 cm wide, erect to slightly nodding, with 2 spikelets per node; internodes 
(5-)7-15(-22) mm, about 0.1-0.3 mm thick at the thinnest sections, with slight 
dorsal ridges, with green longitudinal bands along the concave sides, glabrous 
except for the ciliolate margins; disarticulation below each floret, but not be- 
low the glumes. Spikelets appressed, 25-40 mm (excluding awns), with 5-8 flo- 
rets (including terminal rudiment). Glumes subequal, 14-24 mm long includ- 
ing the undifferentiated awns, 0.1-0.3 mm wide, setaceous, with O-1 distinct 
vein, glabrous, awns more or less straight; lemmas 8-12 mm, glabrous, awns 8- 
25mm, straight, flexuous or slightly curving; paleas about 7-11 mm, obtuse or 
narrowly truncate; anthers 4.5-6 mm, evident in May. 2n = unknown. 


CAMPBELL A 489 


Elymus texensis is known only from three collections that were made on 
calcareous bluffs and hills, in juniper woods and grassy areas on the Edwards 
Plateau of southwest Texas (Campbell 2002). It is similar to E. pringlei, but dif- 
fers in its larger anthers: its glabrous lemmas; its lar ger spikelets; its longer spikes, 
with rachis internodes that are longer, with green longitudinal bands, and gen- 
erally glabrous; and its foliage, which is sometimes densely pilose. It is notable 
that the northernmost known collection of E. pringlei appears somewhat atypi- 
cal and may be transitional to E. interruptus or E. texensis: MEXICO, Coahuila, 
54 mi SE of Big Bend National Park, south end of Sierra Maderas del Carmen, 
Canyon de la Fronteriza, ryolite area, J. Henrickson 15045 (TEX). That collec- 
tion has relatively robust glumes, long rachis internodes, spikelets with up to 6 
florets (including the terminal rudiment), and lemmas that are virtually gla- 
brous (Campbell 2002a). This new species is named after the great state of Texas. 


ParATYPES: TEXAS. Burnet Co.: Inks Lake State Park, limestone bluffs, juniper woods along creek just 
east of HQ, 19 May 1983, R. & G. Kral 70066 (VDB, SMU-BRIT). Uvalde Co.: chalk bluff on Nueces 
River, 12 May 1938, V.L. Cory 29073 (US 3039432). 


KEY TO-FLYMUS FYS ER EAGCAND ITS: ALLIES 


These two new species appear to belong ina natural group with Elymus hystrix 
and other allies. The following key distinguishes the members of this group. 
Brief notes on ranges are inserted, but Campbell (2002a) should be consulted 
for maps and further details. 

This group of species is characterized by their reduced or vestigial glumes, 
with O-1(-2) veins, tapering from the base, 0-24 mm long including awns, of - 
ten differing in length by at least 4 mm, 0.1-0.5(-0.7) mm wide, persistent after 
florets disarticulate; and by their rachis internodes usually 0.1-0.3 mm thick at 
the thinnest sections, often with green longitudinal bands along the concave 
sides. This group should be contrasted with the more heterogeneous group that 
consists of E. virginicus L., E. canadensis L., E. glaucus Buckley and their allies. 
The latter species have glumes with 2-5(-8) veins, widening or linear above the 
base, 4-45 mm long including awns, subequal, 0.2-2.3 mm wide, persistent or 
disarticulating: and rachis internodes usually 0.2-0.8 mm thick at the thinnest 
sections, usually lacking green bands. Elymus interruptus remains a somewhat 
problematic taxon, with specimens that can fit within either of these outlines. 
Moreover, various hybrids of E. hystrix with other species will fit within either 
outline, and will require deeper analysis. It is likely that some allies of E. hystrix 
in this group originated from Peeauar (Church 1967; ee 2002a). 


1. Spikelets widely divergent to perpendicular at traight (rarely 
slightly curving); glumes vestigial or 1-3 mm long, ea some ee 
glumes up to 10(-20) mm long and 0.1-0.2 mm wide but with no distinct vein; 


spikes more or less erect [widespread in eastern North America, but unknown in 
Texas] E. hystrix 


490 BRIT.ORG/SIDA 22(1 


Sie 
Se 
Soa a 


L 
— 


Fic. 2a Fly is(2ad fi E.S. Nixon 537). A. Habit. B Upy I ti f cul it pil , Viewed on 
1 ey I j £ 1 L « ILI J L ry) a Tl rf £. 2 J 
hairs EF AAA ee J | 1 _ ans 1 £ *1 | rs “aL L oe * £ = 
~ i F Fr t J 
spikelet, and | ly si ). F. Spikelet. with lateral vi f fl G. Mature fl 


(left) and adaxial view (right). H. Cross-sections of mature, indurate gl base(s). I. Cross-section of central rachis 
internode. 


a= 


CAMPBELL, A 491 


Fic. 2b. Ely is(2bd fi VL. Cory 29073). A. Habit. B. Upper porti f cul it t pil viewed on 


f g I | D Al £ _ g 

hairs E.M hisi | 1gl ; i di pl f pil | I i g 
spikelet, and | gely sid Igl ). F. Spikelet, with | lvi f fl G. Mature floret in abaxial vi 
(left) and adaxial view (right). H. Cross-sections of mature, indurate gl base(s). I. Cross-section of central rachis 
internode. 


492 BRIT.ORG/SIDA 22(1) 


1. Spikelets usually appressed, never perpendicular; lemma awns straight or curving; 
glumes sometimes vestigial, but usually 1-24 mm long, 0.1-0.5(-0.7) mm wide, 
often with a distinct vein; spikes erect, nodding or penden 
2. ‘ ic well-developed, at least 12 mm long, Sieetel lemma awns straight 

moderately curving; spikes erect to slightly nodding. 
aes (6-)9-15(—22) mm long (excluding awns), each with 2-5 florets; 
lemma awns moderately outcurving at maturity; glumes 0.2-0.5(-0.7) mm 
wide [in the southern Rocky Mountains and adjacent regions, from southern 
California to ea Texas to northern Coahuila E. interruptus 
3. Spikelets 18-40 mm long (excluding awns), each with 3-8 florets; lemma awns 
straight to a curving at maturity; glumes 0.1-0.3(-0.6) mm 
4, Anthers 2.5-4 mm long;lemmas eae hispid to thinly strigose soils 
cent; spikelets 18-25 mm long, with 3-6 florets; spikes 4-12 cm long, ra- 
chis internodes 3-6 mm lo eter ee longitudinal bands, hispid on 
dorsal ridges; blades thinly scabridulous, hispidulous or pilose on veins 
[along the Sierra Madre Orientale, in eastern Mexico from Coahuila to 
Veracruz] E. pringlei 
4. Anthers 4.5-6 mm long;lemmas glabrous; spikelets 25-40 mm long, with 
sieaa alee ae cm lenig co internodes (5-)7-15(-22) mm long, 
pt for ciliolate margins; blades 
thinly scabrous -hirsutet to dense ely pilose [on the Edwards Plateau, in south 


Tex E. texensis 
2. Some glumes less than 12 ae pair usually differing in length by at least 4 
mm when developed, or one or both vestigial: lemma awns [eon at ma- 
a often oe SO; ve aaa nodding to penden 
5. Rachis inter —6(-9) mm long;glumes (0.1-)0.2—-0.5(—0.6) mm wide; lem 


mas eee to strigose, at least near margins; sheaths aeane plants not 
glaucous to moderately glaucous [mostly in the northern Great Plains, from 
Saskatchewan to Ontario to lowa] E. diversiglumis 
. Rachis internodes (4—)6-13(-18) mm long; glumes 0.1—-0.3 mm wide; lemmas 
glabrous or pubescent; sheaths glabrous or villous; plants usually glaucous, 
sometimes strongly so 
6b usually pubescent, the awns (10-)20-30(-35) mm long; spikelets 
with 3(5) florets; rachis internodes (5-)7—13(-—18) mm long, with green lon- 
gitudinal bands and hispid dorsal ridges; blades glabrous or short-pilose; 
plants not strongly glaucous [central Ouachita and western Ozark moun- 
ns in Arkansas, Missouri and Oklahoma] E. churchii 
glabrous, or occasionally arate near apex, the awns (8-)10—20 
(-25) am ng; spikelets with (3—)4—5 florets; oo ~)6-10(-12) 
mm long, without green longitudinal bands, glabrous; blades usually vil- 
lous; plants strongly glaucous [central Interior Low Plateaus, in Kentucky 
and Tennessee] E. svensonii 


n 


OTHER NOTES 


The taxonomy of Elymusin North America hasa long tortuous history. In draft- 
ing a treatment of the species with paired spikelets, | have made some sugges- 
tions for nomenclature, which will hopefully guide botanists in their interpre- 
tation of these taxa (Campbell 1995, 1996, 2000, 2002a, 2002b, 2002c). However, 
it is clear that much deeper analysis is needed, including a proper focus at the 


CAMPBELL A 493 


genetic level, instead of relying just on the rather inadequate morphological 
differences. There has been much taxonomic confusion, misidentification, and 
erroneous reporting of distributions. Many mysteries remain. 


An Elymus hystrix-like collection from New Mexico 

Ananomalous collection that may have similarities to both Elymus hystrix and 
E. interruptus is of interest here: NEW MEXICO [Colfax Co.], Cimarron, wooded 
bank, 6 Jul 1939, W.A. Silveus 4928 (TEX). This specimen was listed by Church 
(1967) under his “atypical” E. hystrix group with filiform glumes. Unfortunately 
all the florets are lost, but the glumes are narrower (ca. 0.2 mm) and more widely 
spreading than typical E. interruptus, and rachis internodes are shorter (ca. 5 
mm). This specimen suggests that plants closely related to E. hystrix may still 
be found in northern New Mexico or nearby. The closest documented E. hystrix 
is in eastern Oklahoma. Does this New Mexico plant have a distinct origin, per- 
haps from some isolated introgressed population? 


Elymus interruptus in California 

[recently discovered the following specimen of Elymus interruptus: CALIFOR- 
NIA, Fresno Co., Pine Ridge, altitude 5300 ft, “Plants of the Sierra Nevada Moun- 
tains,” 15-25 Jul 1900, H.M. Hall & H.P Chandler 317 DOV). This collection was 
initially annotated as E. canadensis. It appears to be the first record of E. inter- 
ruptus from California. The few other records of E. canadensis from the state 
should be checked. 


Need to clarify typification of Elymus virginicus 

Some of the Linnaean material of Elymus virginicus does not clearly match our 
current typical concept of that species, and should probably be included with 
E. glabriflorus (Vasey ex L.H. Dewey) Scribn. & CR. Ball. | proposed to the Inter- 
national Association for Plant Taxonomy that the name E. virginicus be retained 
for the typical plants of Hitchcock and Chase (1951) and most other authors, 
with a new type established for that species (Campbell 1996). However, more 
careful analysis of the several sheets of Linnaean material is needed, and the 
Nomenclature Committee remains undecided (D. Nicholson & R. Soreng, pers. 
comm.). After the treatment in Flora of North America is published, there will 
undoubtedly be further consideration of this problem, and I hope to revive the 
proposal with more evidence. 


Corrections to Campbell (2002b) 

(a) In the key, leads (number 13 & 14) to E. glaucus, E. hirsutus J. Presl. and E. 
dahuricus Turcz. ex Griseb. will be modified substantially for the Flora of North 
America. E.dahuricus isa rare Asian introduction that appears close to E. hirsutus; 
its lemmas can be glabrous, not just scabrous or hispid, as stated in the key. 

(b) In the key, the lead (number 15) to E. interruptus versus E. canadensis and E. 
wiegandii Fernald should omit the spike internode thickness character, which 
reflected some erroneous data. 


494 BRIT.ORG/SIDA 22(1) 


(c) By priority, the correct name for Elymus submuticus (Hook.) Smyth & Smyth 
is E. curvatus Piper. 


ACKNOWLEDGMENTS 


lam deeply grateful to Mary E. Bark worth, who has provided continual encour- 
agement and constructive advice during the 20 years that I have dabbled ec- 
centrically in Elymus. Without her invitation to contribute to the Manual of 
Grasses, and then the Flora of North of America, my efforts would not have 
progressed. She provided insightful comments on an earlier draft of this manu- 
script. [am grateful also to Kathleen M. Capels, who corrected several errors 
and inconsistencies in the key and pointed out the priority of E. curvatus. 


REFERENCES 


Campsett, JJ.N. 1995. New combinations in eastern North American Elymus (Poaceae). 
Novon 5:128. 

Campeett, J.J.N. 1996. Proposal to conserve Elymus virginicus (Poaceae) with a conserved 
type. Taxon 45:128-129. 

Campeett, J.J.N.2000.Notes on North American Elymus species (Poaceae) with paired spike- 
lets. |.E. macgregorii sp.nov.and E.glaucus ssp. mackenzii comb. nov. J. Kentucky Acad. 
Sci. 61:88-98. 

Campeett, JJ.N. 2002a. Notes on North American Elymus species (Poaceae) with paired 
spikelets. Il. The interruptus group. J. Kentucky Acad. Sci. 63:19-38. 

Campsett, J.J.N. 2002b, Notes on North American Elymus species (Poaceae) with paired 
spikelets. Ill. A synoptic key. J. Kentucky Acad. Sci. 63:39-46. 

Camesett, J.J.N. 2002c. Notes on North American Elymus species (Poaceae) with paired 
spikelets. IV. A key to the species and varieties in Kentucky. J. Kentucky Acad. Sci. 63: 
47-52. 

Cuurcn, G.L. 1967. Taxonomic and genetic relationships of eastern North American spe- 
cies of Elymus with setaceous glumes. Rhodora 69:121-162. 

HitcHcock, A.S., and A. CHase. 1951, Manual of the grasses of the United States. 2nd ed. 
U.S.D.A. Misc. Pub. 200. 

Smith, E.B. 1991. An atlas and annotated list of the vascular plants of Arkansas. 2nd ed. 
Published by the author, University of Arkansas, Fayetteville. 

STeveRMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames. 


A NEW HYBRID GENUS AND 12 NEW COMBINATIONS 
IN NORTH AMERICAN GRASSES 
Mary E. Barkworth 


Intermountain Herbarium 


tah S 
Logan, Utah 84322-5305, U.S.A. 
mary@bi are 


ABSTRACT 


One new hybrid genus, x Pascoelymus = Pascopyrum X Elymus, and 12 new combinations are pre- 

sented for North American grasses. Six of the new combinations are for hybrids in the Das 

Stipa arnowiae is transferred to Achnatherum and Hystrix californica (= Elymus ) to 

edie sg Egret is aan as a subspecies of Elymus ceoiaiue Avi mophila 
Jon 


n the findings of others, and the previously recog- 


nea subspecies of Peeudormenneria spicata are reduced to forms. Reasons for the changes are 
provided. 


RESUMEN 


Se Diecut un nuevo género hibrido, sbascoelymiis = = mascop uD x A oF V2 ee Snes 
ticede. 
Stipa arnowiae se transfiere a Achnatherum y Hystrix californica (=Elymus cries a ae 


Agropyron riparium se reconoce como una subespecie de Elymus lanceolatus, Ammophila 


s 
= oS 


champlainensis se reduce a una subespecie, basandonos en los hallazgos de otros, y la subespecie 
| 7 eal. p ] } } J £ ie ; mT 
I 


i f 
los cambios. 


INTRODUCTION 


Volume 24 in the Flora of North America series will be the second of the two 
volumes on grasses. Its completion requires publishing the following 12 names. 
Eleven of the 12 new combinations are for previously recognized taxa; one is a 
new hybrid genus. 

Six of the new combinations, and the new genus, are for hybrids in the 
Triticeae. Identifying the parents of lees without knowing which species 
were present in the vicinity is al fficult. In most instances, I have accepted 
the parentage suggested by previous workers when naming these taxa; in one 
instance, I have felt compelled to disagree. The known distribution of many of 
these hybrids is limited. The remaining six names affect non-hybrid taxa. Two 
reflect a generic change and four a change in rank. 

I have attempted to examine type material of all the taxa treated. As indi- 
cated below, however, this has not always been possible. All the taxa mentioned 
will be described and illustrated in the Flora of North America volume 24. The 


SIDA 22(1): 495 — 501. 2006 


= 


496 BRIT.ORG/SIDA 22(1) 


illustrations will also be available at hetp://herbarium.usu.edu/webmanual/. 
To the extent permitted by the institutions that own them, photographic im- 
ages of the type specimens examined are also available via this site and will be 
available via TROPICOS. The images include close ups of individual parts in 
addition to images of the whole specimen. 


NOMENCLATURAL TREATMENT 


POEAE 


Ammophila breviligulata subsp. champlainensis (F Seym.) Walker, Paris & 
Barrington ex Barkworth, comb. nov. Basionym: Ammophila champlainensis F 
Seym., Sida 2:349. 1966. TyPF: U.S.A. NEW York: Lake Cham plain, Au Sable Point, in sand, 3 Jul 
1902, Nellie F. Flynn s.n. (HOLOTYPE: VT). 


— 


Walker, Paris, and Barrington (1998) reported that Ammophila breviligulata 
sensu stricto Fern.and A. champlainensis differ completely in glume length and 
flowering time, and tend to differ in inflorescence length. They noted, however, 
that the morphological and molecular uniformity of A. champlainensis, com- 
bined with the prevalence of vegetative reproduction in both taxa, is consis- 
tent with the hypothesis that all populations of A. champlainensis are derived 
from a single genetic individual. For this reason, they recommended acknowl- 
edging the distinction between the two taxa at the subspecific, rather than spe- 
cific level. 1 present the new combination here so that it may be used in volume 
24 of the Flora of North America. 


= 


STIPEAE 


rece arnowiae a L. Welsh @ N.D. Atwood) Barkworth, comb. nov. 

BASION S.L. Welsh & N.D. Atwood, Utah FI. (ed. 3):799. 2003. U.S.A. UTAH. 
Kane Co. T4+ 35, R4W,S 13,ca 19 mi Eof Johnson Canyon Jct; pinyot liper-sagebrush-com 
munity at 1740 m, on white, gypsiferous member of the vane For mation, 30 May 2001, 
S.L. Welsh & T. O’Dell 28062 (HOLOTYPE: BRY; ISOTYPE UTC-245001). 


= 


This taxon is very similar to Achnatherum hymenoides (Roem. & Schult.) 

Bark worth, differing in having loosely contracted panicles with non-divaricate 

branches. 

TRITICEAE 

<Elyleymus hultenii (Melderis ex Hultén) Barkworth, comb. nov, BasionyM: 
x Agroelymus hultenii Melderis ex Hultén, Ark. Bot, ns. 7(1):2L. 1968. Agropyron alaskanum 
var. arcticum Hultén, Acta Univ. Lund, nus. 38: 257. 1942. Type: U.S.A. ALASKA: Deering, J.P. 
Anderson 4790 (HOLOTYPE: ALA). 

Hultén (1968) listed the parents of this hybrid as Agropyron boreale subsp. 

alaskanus(Scribn. & Merr) Melderis and Elymus arenarius subsp. mollis (Trin.) 

Hulten. The new combination is needed because these two species will be treated 

as FE. alaskanus (Scribn. & Merr.) A. Love subsp. alaskanus and Leymus mollis 

(Trin.) Pilger, respectively, in the Flora of North America vol. 24 


BARKWORTH 497 


Although Hultén (1968) listed Melderis as the author of the name 

x Agroelymus hultenii, he did not explain Melderis’ contribution in the article. 

For this reason, the authorship is “Melderis ex Hultén” rather than “Melderis in 
Hultén”. If using short form citations, it would be Hultén 

<Elyleymus guiness (Lepage) em comb. nov. BAsIONyM: x Agroelymus 

ontariensis Lepage, Nat. Can. 79:254-257. 1952. Type: CANADA. Ontario: James Bay, Riv. 

Attawapiskat, 21 oe 1946, A. Dutilly é & — Lepage 16,423 (HOLOTYPE: originally deposited in 

LCU, transferred to NA, then to US. Despite a search at both NA and US, it has not been lo- 


<Elyleymus ontariensis, according to Bowden (1967), comprises hybrids be- 
tween Elymus trachycaulus (Link) Gould ex Shinners[= Agropyron trachycaulum 
(Link) Malte ex H.E Lewis] and L. innovatus (Beal) Pilger [= Elymus innovatus 
Beal]. Bowden regarded it as a synonym of XElyleymus hirtiflorus (Hitchc.) 
Barkworth & D.R. Dewey, but the holotype of that hybrid grows outside the 
range of L. innovatus. It is interpreted as having Leymus simplex (Scribn. & T.A. 
Williams) D.R. Dewey as the Leymus parent. 


XElyleymus mossii (Lepage) Barkworth, comb. nov. BasionyM: x Agroelymus mossii 
Lepage, Nat. Canad. 92:214-215. 1965. TyPpE: CANADA. mois Near Lake Louise, in open 
woods, 22 Aug 1946, E. H. Moss 7257 (HOLOTYPE: ALTA 10 

The proposed combination reflects a difference of ee concerning the par- 

ents of this hybrid as well asa difference in generic interpretation. Lepage (1965) 

stated that it was obvious that Elymus canadensis L. was one parent of this hy- 

brid, but that it would be necessary to discover which species of Agropyron lin 
the traditional sense] grew in the neighborhood to determine the other parent. 

He gave “Agropyron (?) trachycaulum’ as a possibility. Boivin (1967) agreed that 

E. canadensis but suggested that the Agropyron parent was Agropyron violaceum 

(Hornem.) Lange [= Elymus violaceus (Hornem.) Feilberg]. 

Elymus canadensis, however, is generally absent from the region around 
Lake Louise (Moss 1983), the area where the holotype was collected. Other fea- 
tures of the holotype that are difficult to reconcile with Lepage’s suggested par- 
entage are the presence of rhizomes and the abundance of relatively long soft 
hairs on both the lemmas and the glumes. Elymus canadensis is not rhizoma- 
tous, has coarse hairs on the lemmas, and has scabrous rather than hairy glumes. 

Elymus trachycaulus and its high elevation counterpart, E. alaskanus subsp. 
latiglumis(Scribn. &J.G. Sm.) A. Léve, usually lack rhizomes, and have glabrous 
or shortly hairy lemmas and glumes. More probable parents are Elymus glaucus 

Buckley and Leymus innovatus, both of which are common in the region. Elymus 

glaucus has spikes that more closely resemble the holotype in their posture, 

thickness, and awn length than those of E. canadensis; L. innovatus has soft hairs 
on its glumes and lemmas. Elymus glaucus is sometimes shortly rhizomatous; 

Leymus innovatus is always rhizomatous. Based on this interpretation of the 

parentage, the hybrid has to be included in x Elyleymus. 


498 BRIT.ORG/SIDA 22(1) 


Elymus Xcayouetteorum (Boivin) Barkworth, comb. nov. BasionyM: x Agrohordeum 
cayouetteorum Boivin, Nat. Canad. 94:520. 1967. TYPE: QUE! 

Boivin (1967) published this name for hybrids between Elymus trachycaulus[= 

Agropyron trachycaulum] and E. canadensis. 

Elymus lanceolatus subsp. riparius (Scribn. & J.G. Smith) Barkworth, comb. & 
Stat. NOV. BASIONYM: Agropyron riparium Scribn. & J.G. Smith, Bull. Div. Agrostol., U.S.D.A. 
4:35. 1897. TyPE: U.S.A. MONTANA: Garrison, 10 Jul 1895, PA. Rydberg 2127 (LECTOTYPE: US 
556672), designated by Hitchcock 1935:776). 


After examining many specimens, I agree with Dorn (1988) that this taxon 

merits recognition but prefer to treat it as a subspecies rather than a variety. It 

is more common than Elymus lanceolatus (Scribn. & J.G. Sm.) Gould subsp. 
lanceolatus. 

x Leydeum littorale (HJ. Hodgs. @ W.W. Mitch.) Barkworth, comb. nov. BAsIonYM: 
xElymordeum littorale HJ. Hodgs. & W.W. Mitch., Canad. J. Bot. 43:1355. 1965. TyPE: U.S.A. 
ALASKA: Matanuska Valley, in tidal flat area along Cottonwood Creek near juncture with Knik 
Arm about 18 mi SW of Palmer, 2 Sep 1964, W.W. Mitchell & HJ. Hodgson 1584 (HOLOTYPE: 
ALA 29247)). 


md. 


xLeydeum littorale consists of hybrids between Leymus mollis and Hordeum 
brachyantherum Nevski. It has been collected in the Matanuska Valley, Alaska, 
and on the coast of Vancouver Island, British Columbia and it may be more 
widespread. Hodgson and Mitchell (1965) stated that it grows along the mar- 
gins of the high-tide zone, where tidal waters flow into the creek. The hybrid 
plants can easily be distinguished from L. mollis by their shorter narrower spikes 
and more yellow and dense foliage. The new combination reflects recognition 
of the segregate genus Leymus. 

Leymus californicus (Bol. ex Thurber) Barkworth, comb. nov. BasionyM: 
Gymnostichum californicum Bol. ex Thurber in S. Watson, Bot. California 2:327. 1880. 
Gymnostichum californicum Bol. ex Thurb., Bot. California 2:327. 1880. Hystrix californica 
(Bol. ex Thurb.) Kuntze, Revis. Gen. Pl. 2:778. 1891. Type: Elymus californicus (Bol. ex a 
Gould, Madrono 9(4):127. 1947. U.S.A. CALIFORNIA: Redwoods, near San Francisco, H.N. 
Bolander s.n. (LECTOTYPE: GH-19493), designated by Baden, Frederiksen & Seberg, Nordic J. 
Bot. 17:457. 1997). 


Transfer of this taxon to Leymus is supported by its chromosome number of 2n 
= 56, and genome-specific RAPD assay results Jensen & Wang 1997). It is con- 
sistent with Mason-Gamer’s (2001) examination of granule-bound starch syn- 
thase genes in allotetraploid Triticeae. In her tree, it groups with Psathyrostachys, 
Nevski, a genus very close to Leymus (Bodvarsdottir and Anamthawat-Jonsson 
2003). Morphological characteristics that tend to place it in Leymus rather than 
Elymus include its well-developed rhizomes and the more or less equally promi- 
nent veins in its leaf blades. There are also many specimens of L. innovatus, 
particularly those from Alaska and Yukon Territory, that lack or have very re- 
uced glumes. 


BARKWORTH, NEW NAMES 499 


The habitat of L. californicus, coniferous forests on non-alkaline soils, is 
unusual among North American species of Leymus. It is, however, similar to 
that of some Chinese species that are currently included in Hystrix Moench 
because of their lack of glumes. 

Hitchcock (1935, 1951) and Baden et al. (1997) included Leymus californicus 
in Hystrix because it lacks glumes. The type of Hystrix is, however, Elymus 
hystrix, a species that, apart from lacking glumes and having strongly diver- 
gent spikelets, is mort lly, genomically, and molecularly similar to other 
eastern North American species of Elymus. 


<Pascoleymus Barkworth, gen. hybr. nov. = shane a A. Love x Leymus 
Hochst. Tyee: Pascoleymus bowdenii (Boivin) Barkwor 
XPascoleymus bowdenii (Boivin) Barkworth, a NOV. BASIONYM: X Agroelymus 
owdenii Boivin. Nat. Canad. 94:520. 1967. Type: CANADA. ALBERTA: Beaverlodge, 19 Jul 1921, 
M.O. Malte 108211 (HOLOTYPE: QFA!). 


< Agroelymus bowdenii applies to hybrids between Agropyron smithii Rydb. 
[=Pascopyrum smithii (Rydb.) A. Love] and Elymus innovatus [= Leymus 
innovatus] (Boivin 1967). Recognition of the genus Pascopyrum requires a new 
intergeneric hybrid name as well as a new combination at the species level. 


Pseudoroegneria spicata [. inermis (Scribn. & J.G. Sm.) Bark worth, comb. nov. & 

Stat. NOV. BASIONYM: a divergens var. inerme Scribner &J.G.Sm., Bull. Div. Agrostol 

US.D.A. 4:27. 1897. TYPE: U.S.A. IDAHO: 1895, L.FE Henderson 3058 (LECTOTYPE: US-556676), 
Hitchcock 1935:773). 


Daubenmire (1939, 1960) reported that rhizome development and awn length 
in Agropyron spicatum (Pursh) Scribn. & J.G. Sm. [= Pseudoregoegneria spicata 
(Pursh) A. Love] varied continuously within plants grown from seed. He con- 
cluded that the ability to produce rhizomes and unawned plants is heritable, 
that the two characters are not linked, and that which form becomes dominant 
at a local site is determined by environmental conditions. 

Plant breeders working with Pseudoroegneria spicata consider that awn 
presence is determined by a single major gene, modified by some minor genes, 
with the unawned condition being dominant. Although no one has gathered 
the data that explicitly tests this hypothesis, extensive work with both awned 
and unawned accessions of the species suggests that it is true. It means that a 
pair of heterozygotic unawned parents will give rise to around 50% awned off- 
spring. This hypothesis is consistent with Daubenmire’s observations. 

The above observations make it clear that the awned (P spicata f. spicata) 
and unawned (P spicata f. inermis) phases of Pseudoroegneria spicata are of little 
taxonomic significance despite their evident morphological difference. The rea- 
son for making names available at the level of form is to accommodate those 
who wish to distinguish the two entities. 


P ia spicata f. pubescens (Elmer) Barkworth, comb. nov. & stat. nov. 


So 


BRIT.ORG/SIDA 22(1) 


BASIONYM: Agr Py spicatum var. pubescens Elmer, Bot. Gaz. 36:52. 1903. Agropyron spicatum 

subsp. puberulentum Piper, Contr. U.S. Natl. Herb. 11:147.1906, nom. superfl. TYPE: U.S.A. WASH- 
INGTON: Kittitas Co. Mt. Stuart, Jul 1898, A.D.E. Elmer 1158 (HOLOTYPE: Deposited in Stanford 
Herbarium which has been transferred to CAS; despite a search, it could not be located at 


CAS: ISOTYPE: US-1817092!) 


Plants of Pseudoroegneria spicata with densely pubescent leaves are known 
from the east slope of the Cascade Mountains in Washington. Because plants 
with nearly as densely pubescent leaves are found elsewhere in southern Wash- 
ington and northeastern Oregon, it seems best to recognize them as a form, P. 
spicata f. pubescens, commensurate in this respect with the level of recognition 
given the awned and unawned phases. 

Publication of the above combinations automatically generates 
Pseudoroegneria spicata (Pursh) A. Love f. spicata with priority dating from 1814, 
the year of publication of the basionym for the specific epithet. 

ACKNOWLEDGMENTS 
Ithank the curators of the herbaria from which | have borrowed the specimens 
cited in this paper, and those at other herbaria who have assisted me in locating 
the types that have been transferred from their original institution. also thank 
Kanchi Gandhi for clarifying some of the nomenclatural issues involved, James 
Zarucchi, and Kathleen Capels for assistance in obtaining copies of the relevant 
literature, and Paul Peterson for his prompt and helpful review. 


REFERENCES 


Baoen, C., S. FREDERIKSEN, and O. SeBerc.1997. A taxonomic revision of the genus Hystrix 
(Triticeae, Poaceae). Nordic J. Bot. 17:449-467, 
BODvARSDOTTIR, S.K.and K. ANAMTHAWAT-JONSSON. 2003. Isolation, characterization, and analysis 


of Leymus-specific DNA sequences. Genome 46:673-682. 

Boivin, B. 1967. Enumérations des plantes du Canada VI—Monopsides, (2eme partie). 
Naturaliste Canad. 94:471-528. 

Bowpen, W.M. 1967. Taxonomy of intergeneric hybrids of the tribe Triticeae from North 
America. Canad. J. Bot.45:711-724. 

Dausenmire, R. 1939. The taxonomy and ecology of Agropyron spicatum and A. inerme. Bull. 
Torrey Bot. Club 66:327-329. 

Dausenmire, R. 1960. An experimental study of variation in the Agropyron spicatum—A. 
inerme complex. Bot. Gaz. 122:104-108. 

Dorn, R.D. 1988. Vascular plants of Wyoming. Mountain West Publishing, Cheyenne, Wyo- 


~— 


ming. 340 pp. 

HitcHcock, A.S. 1935. Manual of grasses of the United States. U.S. Government Printing 
Office, Washington, D.C 

HitcHcock, A.S. 1951.Manual of grasses of the United States, ed. 2, revised by A.Chase. Misc. 
Publ. 200. U.S.Government Printing Office, Washington, D.C 


BARKWORTH, NEW NAME 501 


Hopcson, H.J.and W.W. MitcHet. 1965.A new Elymordeum hybrid from Alaska. Canad. J.Bot. 
43:1355-1358. 

Hutten, E. 1968.Comments on the Flora of Alaska and Yukon. Ark. Bot. 7:1-147, 

Jensen, K.B. and R.R.-C. Wane. 1997. Cytological and molecular evidence for transferring 
Elymus coreanus from the genus Elymus to Leymus and molecular evidence for Elymus 
californicus (Poaceae: Triticeae). Int. J. Plant. Sci. 158:872—877. 

Lerace, E. 1965. Révision généalogique de quelques XAgroleymus. Naturaliste Canad. 
92:203-216. 

Mason-Gamer, R.J. 2001. Origin of North American Elymus (Poaceae: Triticeae) Allotetrap- 
loids based on granule-bound starch synthase gene sequences. Syst. Bot. 26:75 7-768. 

Moss, E.H.1983.Flora of Alberta, ed. 2, revised by J.G. Packer. Univ.of Toronto Press, Toronto, 
Ontario. 

Waxker, PJ., CA. Paris, and D.S. Barrincton. 1998. Taxonomy and phylogeography of the 
North American beachgrasses. Amer. J. Bot. 85:87. 


502 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


ANN Linpsay and Syp House. 2005. The Tree Collector: The Life and Explorations 
of David Douglas. (Paperback Edition). (ISBN 1-84513-052-9, pbk.). Aurum 
Press, 25 Bedford Ave., London WCIB 3AT, UK, distributed by Trafalgar Square, 
(Orders: 802-457-1911, 802-457-1913 fax, www.trafalgarsquarebooks.com). 
$16.00, 256 pp., an bibliography, index, b&w photos, figures and 
maps, 5" x 7 3/4" 


The book The Tree Collector by Lindsay and H ul | i h it and 
adventures of botanical expeditions of David Douglas, the Scotsman foe which the Douglas fir tree 
was named. The authors have woven a wonderful story of the botanical work of David Douglas by 
interlacing history and background information with David Douglas's own journal excerpts. Lind- 

say and House's efforts have created a book in which one really feels as though they are along for the 


journe 

eee en expeditions for the Horticultural S occur alter Lewis and Clark's jour- 
neys (1804-1806) and before any major Indian Wars (1840). Douglas's own words an ad journal en- 
tries are the foundation of the story. Readers get wonderful descriptions of Douglas's impressions of 
plants, animals, |: ape and climate in the Eastern United States, Canada, Pacific Northwest, Cali- 
fornia and Hawaii as well as insightful information on the indigenous peoples he encounters along 
his expeditions. Douglas's journal entries also include hardships with weather, illnesses, ane hs 
and disappointments in his collections and the outcome of his return to ae onder, which make this 
book all the more real for the reader. Truly engaging are stories of D itement in Gatie 
down plants of his particular interest such as the sugar pine. 

The book includes an introduction of the family history of David Douglas and how he came to 

be a member of the Horticultural society the ero that oe ted ee se ceemans Great c 
merica aon 


° 
= 


is taken in providing details of 
eka eee with Dovsee journal entries, eae and white line maps ae images are included to 


themselves with locations discussed. Appendices included in the text are excel- 

lent summaries of information on Dons’ eae uaa to science, such as species in- 

troduced and those named after g pithet are included). as arenes. 
ilsin Scotland, North Aspens aand Hawaii. | 


that discuss ae Botanical introductions include short descriptions of the plant from 
ouglas’s journal, as well as the authors’ information on ms or other factual information 
Take a journey with David Douglas into the wilds of North America and discover new plants, 
animals, and peoples... pick up a copy of The Tree ia today!—Lee Luckeydoo, Herbarium, Bo- 
tanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(1): 502. 2006 


QUADRUPLE, TRIPLE, DOUBLE, AND SIMPLE PAPPI IN THE 
GOLDENASTERS, SUBTRIBE CHRYSOPSIDINAE 
(ASTERACEAE: ASTEREAE) 


John C.Semple 


Depar rtment of Biology 
e 


jcsemple@sciborg.uwaterloo.ca 


ABSTRACT 


Pappus variation in eight genera, 58 species, and 94 taxa of the Chrysopsidinae was examined; qua- 
druple, triple, double ance sometimes sssimple pape! were ound wi variation in the size, shape and 
les. In the genera witha hehe: 


as of sk 


I 9)—Heterotheca, Noticastrum, Pityopsis,and probably Iomentaurum— 


Ba iene are more numerous and in a greater number of series. This is interpreted as the 
milar to, but with less clavate inner bristles, than found in Doellingeria and 
ae the ae genera in the North American Clade of the Astereae. All four genera have a 


quadruple pappus consisting of four. ling, series: a short dary outer series 
of scales or scaly bristles; a silanes inner series of ae length tapered bristles, a primary outer 
series of tapering bristles 80-95 length of the inner inner series of usually 
weakly clavate bristles. Both ray and disc fl ilar au ie canees ee for the epappose 
ray fruits of H. sect. Heterotheca. 1 ies, the four seri others they grade 


into each other. In Croptilon, the somber of pappus series decreases with the pee reduction 
from x =7 to x = 4. In the lower base number genera Brad buria, Chrysopsis, and Osbertia, a reduction 
in the number of series was observed, with quadruple, ne, ve ane sometimes simple pappi 
occurring. In Chrysopsis, the secondary outer series is well c ped, b inner series 
is absent or reduced to a few bristles. In Osbertia, the pappus Spear nn or includes a second 
series of 1- ; short bristles; all the long bristles are tapering or at best only very weakly clavate. In 
small sample per species, the number of bristles per fruit ranged from 20-125: Noticastrum, 46-125 
ee . ae Tate 80-85; Heterotheca, 23-80 averaging 43 bristles; Pityopsis, 25- 
46, averaging 37; Bradburia, ray florets, 30-36, discs 0-5; Chrysopsis and Croptilon, 18-36 averaging 
28; Osbertia, 20-35, averaging 27. The fruits with lower numbers of bristles had 0-few mid-length 
secondary inner series bristles. 


RESUMEN 


Se examino a variacion e valane en ocho géneros, 58 especies, y ae taxa ace She pslainee se 
I > 


encontraron vi I triples doblesya p forma 


} 1 ] ] ie 


basi As alto (x = 9)—Heterotheca, Noticastrum "Pipi y problemen 


+] 


Tome ntaurum—los sleue ates de) 
se una n primitiva, similar pero con menos seas! interiores ae dé le 


] 


que se ae en nee y Eucephalus 
Astereae. Los cuatro géneros tienen un vilano cuadruple que consta de cuatro series, a veces 
. 1) : fa [ees | q . i ee 


SIDA 22(1): 503 — 531. 2006 


504 BRIT.ORG/SIDA 22(1) 


de sedas afiladas de longitud media; una serie primaria externa de sedas afiladas del 80-95% de la 


longitud de la serie interna; y una serie primaria interna de sedas normalmente clavadas. Tanto las 


flores radiadas como las del disco tienen una ordenacion similar, excepto los frutos sin vilano de H. 


sect. Heterotheca. El alg | t ries son distintas y en otras se intergr oa entre 
ellas. ea | ies del vilano d 


reduccion _ de x = 

En los géneros con numero base inferior Bradburia, pie y Oshertia, se ha ea una 

reduccion en el numero de series, con vilanos cuadruples, triples, do bles /a veces simples. En 

Chrysopsis, la serie secundaria externa esta bien desarrollada, ae : serie secundaria interna esta 

ausente o reducida a unas pocas sedas. En Osbertia, El vil s simple o incluye una cea a serie 
] 


de 1-2 sedas cortas; todas las sedas largas son afiladas 0 com ho solo débi ‘la 


y 


una pequena muestra por especie, el numero de sedas pot fruto vario entre 20-125: Noticastru m, 46- 
125con una media de 87 sedas; eed 80-85; Heterotheca, 23-80 con una see de 43 sedas; 
Pityopsis, 25-46, con una media de 37; Bradburia, flores radiadas, 30-36, del d 5; Chrysopsis 
and Croptilon, 18-36 con una media i: 28; Oshe >rtid, 20-35, con una media de 27. | ae menor 
numero de sedas tuvieron O- 


lew en la serie secundaria interna de longitud media. 
INTRODUCTION 


Goldenaster genera in the subtribe Chrysopsidinae Nesom (Asteraceae: 
Astereae) have long been reported to have a double pappus consisting of a short 
outer whorl of narrow to broad scales or linear bristles (e.g., Gray 1884; Fernald 
1950; Cronquist 1968, 1980; Semple 1981, 1996; Semple & Bowers 1985; Semple et 
al. 1988). Nesom (1994a) in his protologue to subtribe Chrysopsidinae described 
the pappus as “2-3(-4)-seriate, persistent, the inner of 1-2 series of generally 
flattened bristles, outer of much shorter setae, bristles, or scales,” but did not 
elaborate as to the number of series in each genus. Nesom (2000) described the 
pappi of the North American genera as follows: Croptilon, “pappus a single se- 
ries of persistent, thick and rigid, equal-length, tawny to reddish-brown capil- 
lary bristles”; Chrysopsis (including Bradburia), “pappus in 2 series, the outer 

very short bristles, the inner of long capillary bristles”; Heterotheca, “‘pappus of 
tawny to whitish barbellate bristles somewhat uneven in length, witha shorter, 
outer series of lanceolate scales or bristle-like squamellae” and noted the 
epappose to few bristled ray floret cypselae of sect. Heterotheca; Osbertia, “pap- 
pus uniseriate, with numerous, nearly non-barbellate bristles, a short, outer 
series commonly present in O. chihuahuana’, Pityopsis, “pappus 2-seriate, the 
inner of barbellate bristles, the outer of much shorter setiform bristles or se- 
tae”, and Tomentaurum, “pappus of 45-60 white, barbellate bristles in several 
series, with a few inconspicuous setae or very slightly widened bristles, 0.5-L5 
mim long.” Nesom (1994a, 2000) did not indicate which taxa had the flattened 

inner bristles. Noticastrum is native to South America and was not discussed in 

Nesom (2000). Zardini (1985) monographed the genus and described the pap- 

pus as being in two series, the external morphologically similar to the internal 

but of short bristles, normally straw-colored or reddish [five species] to pur- 

plish in N. diffusum and the bristles barbellate especially distally. Zardini also 

noted that the cypsela body was glandular in some taxa. 


fou 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 505 


Many other North American Astereae have been reported to have a simple 
pappus. Some asters often treated in Aster sensu lato have been reported to have 
a “double” pappus (Eucephalus Nutt., Sericocarpus Nees: Gray 1884; Cronquist 
1955) or a “triple” pappus (Doellingeria Nees: Cronquist 1968, 1980; Nesom 
1994a,b; Semple et al. 2002). The “triple” pappus reportedly had two inner 
whorls, the outer slightly shorter and tapering and the inner bristles clavate. 
However, Hood and Semple (2003) demonstrated that nearly all species of gold- 
enrods, which had been reported to have a simple pappus, in fact had a double 
pappus with two primary series of long bristles (the outer shorter and taper- 
ing, the inner clavate) and at least some species had a secondary outer series of 
a few very short bristles, ie. Solidago has a double or triple pappus. Semple and 
Hood (2005) demonstrated that many North American aster genera have a 
double, triple, or quadruple pappus and concluded that the likely primitive 
pappus of the North American Astereae consists of four series: the primary in- 
ner series of long, clavate (flattened) bristles, a slightly shorter primary outer 
series of long tapering bristles,a secondary inner series of tapering bristles 40- 
80% the length of the primary inner bristles, and a secondary outer series of 
short bristles or narrow to broad squamellae usually only 10-15% the length of 
the inner primary bristles. Hood and Semple (2003) proposed a terminology 
for a triple pappus, ie. secondary outer whorl, primary outer whorl, and pri- 
mary inner whorl bristles, which Semple and Hood (2005) modified to cover 
the discovery of a fourth series of bristles found in asters but not seen in gold- 
enrods. They also noted that “series” was a more accurate label than “whorl” 
based on observations of both North American and Old World aster genera. 
They concluded that due to the subtlety of the differences between the longer 
bristle series in many species, it would be practical to treat the pappus of 
Symphyotrichum as “appearing simple” in keys to identification. The details of 
the “double pappus” of the Chrysopsidinae were re-examined in light of the 
recent discoveries on pappus traits of goldenrods and asters and the results are 
presented in this paper. 


MATERIALS AND METHODS 


A preliminary survey was undertaken to examine the pappus of one or two 
specimens of representative species of the genera of the Chrysopsidinae sensu 
Nesom (2000). Subsequently, a more detailed survey was conducted involving 
eight genera, 58 species, and 94 taxa of subtribe Chrysopsidinae listed in Table 
1. Observations were made using a dissecting scope (10-70) or a compound 
light microscope (20-400). The degree of the clavateness of bristle tips was 
determined using the 0-4 rankings described in detail in Hood and Semple 
(2003). At least five different fruits from each species were observed under the 
dissecting microscope at a maximum of 70x; in some cases many fruits from 


ie ] 


many individuals examined to investigate the frequency of variation. The 


506 BRIT.ORG/SIDA 22(1) 


majority of observations were made on specimens in the WAT Herbarium, ad- 
ditional specimens on loan from MEXU, MO, OS, SD, TEX-LL and UC (Holmgren 
et al. 1990) were also ined. In addition to the several methods used in evalu- 
ating pappus features listed by Hood and Semple (2003), assessment of the char- 
acteristics of the short secondary outer scales or bristles was also systemati- 
cally undertaken with three traits recorded. The large difference in numbers of 
pappus bristles per fruit noted in the preliminary survey and the literature led 
to recording a rough estimate of bristle number for each taxon by counting the 
number of bristles on three to five representative fruits per taxon. These were 
compared to counts of numbers of bristles in (Smith 1965; Nesom 1991b; Semple 
& Bowers 1995; Semple 1996). Thus, Table | has eight columns of observational 
data plus a column for additional comments, while only four were reported by 
Hood and Semple (2003) and five in Semple and Hood (2005). 

Observations on the compound microscope were made from both un- 
mounted fruits and from slides prepared following Semple and Hood (2005). 
For each species, usually two to three disc floret cypselae with or without co- 
rollas were mounted in Cytoseal-60'™ mounting medium under a cover slip; 
immature ray floret fruits were also often mounted on the same slide for com- 
parison. Observations at 20-400 on the compound microscope were made 
similarly to the observations under the dissecting scope at 30-70. Observa- 
tions made on the two kinds of scopes were compared and any discrepancies 
were resolved by re-examining specimens. 

Digital photomicrographs were taken using a Nikon CoolPix 990 camera 
manually held against the ocular lens of either the dissecting or compound mi- 
croscope. Pictures were taken of specimens under the compound light micro- 
scope with either below stage or above stage lighting. Scanning Electron Micro- 
scope (SEM) photomicrographs were made using Kodak FX or Ilford Pan F Plus 
film commercially developed and subsequently digitized by scanning the nega- 
tives. Final digital illustrations were made using CorelDraw 12® from digital im- 
ages edited with Corel PhotoPaintl2® (Corel Corp.). In some illustrations, the con- 
trast was manipulated to increase the difference between pappus bristles and 
background for illustrative purposes; backgrounds were darkened considerably 
and sometimes bristles were lightened somewhat to correct for uneven lighting 
and exposure; the converse was done in bristle tip silhouette illustrations 


RESULTS AND DISCUSSION 


The pappus of most species of the Chrysopsidinae consists of 3-4 series (Figs. 
1-7) while a minority of taxa has a pappus of 1-2 series (Fig. 8). The quadruple 
pappus consists of the following series: |) a well develop secondary outer series 
usually of many, short, linear-bristly to broad scales; 2) a secondary inner se- 
ries of mid-length, tapering, non-clavate bristles that were generally 40-80% 
the length of the primary inner series (Figs. LE J, 2F 2N, 4K, 5A, 5M, 6A); 3) a 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 


eile 
onion sone nee 


Fic. 1. Fruit and ts of Heteroth lisc fruit pt A; scal Tmmin A and F,=0.5 mm in C-D and H—- 
| and = 10 min E.A-B.H. 9 liflora (Semple & Semple 5575 WAT). A. Mature epapp yand papr 
i (long) and bristles. CE. H. inuloid ( & Woodland 1928 WAT), 
SEMs. C. Tip of mid lengt ths Jary inner bristle. D. Secondary outer series scales and overlapping bases of longer 
bee peal of barbs on me person of inner series PBUE: F- a H, mexicana Uppne el mall: F. paawe fruit 
( ).G. Upper p \ )and f y ou 
LH. isessiliflara subsp bolanderioides (Semple & Chmiel ki 8918a WAT: SEMs) H Weakl f pri 
series bristles.1.5 lary les.J.H. (H 671WAl), tip pemg length secondary inner iste 
Hee en ee (Semple 9902 WAT), c tips of primary i 
primary outer bristles. L- Lt . oregona var pacta (Semple & Heard 5693 WAT).L. Silhouette of tion of mid 
lengtt 1 I 1 long primary outer and inner bristles. M. Linear secondary auter bristle (arrow). 


TCTIQUIT SCCUNUaTy TIC 


508 BRIT.ORG/SIDA 22(1) 


pH 


primary outer series of tapering bristles that were 85-95% the length of the 
primary inner series (Figs. 1B, 1], lJ, 3D, 4D, 4G, 4K, 5G, 5K, 6KC, 7B, 7E, 71, 7K, 
8A, 8C): and 4)a primary inner series of very subtly to moderately clavate tipped 
bristles (Figs. 1B, HI, 4D, 5B, 0D, 7E, 7J). A four-series pappus is plesiomorphic in 
the North American Clade (sensu Noyes & Rieseburg 1999) of the tribe Astereae 
(Semple & Hood 2005). Heterotheca, Noticastrum, and Pityopsis, with the 
plesiomorphic chromosomal base number of x = 9, has a quadruple pappus in 
all or most species, as does the monotypic Tomentaurum, whose chromosome 
number is not yet known but is hypothesized here to be x=9 based on indument 
and fruit similarities with x = 9 Noticastrum. 

In the genera with lower chromosomal base numbers, Croptilon (synonym: 
Haplopappus sect. Isopappus: Hall 1928), Chrysopsis, and Bradburia, the pap- 
pus is usually reduced to three pappus series or less but sometimes had I-few 
bristles of one or both of the missing series. Osbertia, which also was included 
in Haplopappus by Hall (1928), has a vestigial secondary series and a possible 
biseriate primary series. Thus, in all genera of the Chrysopsidinae, some evi- 
dence was found for the presence of at least 2-3 pappus series and usually more. 

A large range in pappus lengths and numbers of bristles was observed 
among genera and within some genera of Chrysopsidinae. The smallest fruits 
with the lowest numbers of bristles were seen in Bradburia, Chrysopsis, 
Croptilon, and Osbertia. These traits vary considerably in Heterot 
Pityopsis. The majority of species of Noticastrum examined had the longest 
pappus bristles and the highest number of bristles observed. Pappus and fruit 
traits are discussed in detail below by genus with higher base number genera 
presented first. Data for all taxa examined are presented in Table 1. 

Heterotheca (x = 9; all 24 species, all 48 taxa examined).—Cypselae straw 
colored to brown, obconic, compressed laterally or not, those of ray florets of - 
ten 3-angled, 1-4.3 mm long, sparsely to moderately strigose, dimorphic in sect. 
Heterotheca (Fig. 1A), ray florets usually lacking a pappus and glabrous, rarely 
pappose and intermediate; ribs 5-10, sometimes dark, resin filled; pappi qua- 
druple or sometimes triple through loss of the secondary inner series, series 
often intergrading; 23-80 mid to long bristles, averaging about 43 per fruit for 
the genus, barbs usually evenly arranged around the round shaft for most or all 
the length of the bristle; secondary outer series obvious to inconspicuous, of 
linear to narrowly triangular or oblong scales (Figs. LD, I) or linear bristles (Fig. 
IM), these usually 5-15(-30)% the length of the longest primary bristles; rarely 
up to 40% (some specimens of H. mucronata var. harmsiana), secondary inner 
bristles many (H. oregona, Fig. lL) to few or absent (e.g., H. villosa var. nana), 35- 
85% the length of the longest primary bristles, sometimes grading in length 
from linear secondary outer bristles into the primary outer bristles (e.g., H. 
brandegei, H. stenophylla, H. thiniicola), primary inner bristles tapering dis- 
tally, 85-95% the length of the primary inner bristles, fewer than the primary 


— 


heca and 


Taste 1. Pappus variation in subtribe Chrysopsidinae; disc floret cypselae (ray fruit for B hirtella). Clv, degree of clavateness of primary inner bristles (0 = not 
clavate to4= svongy clavate), Clv- -Tap, clavate alternating with tapered bristles (primary inner and outer whorls, y = yes, - not obviously so); Lgth, primary 
outer bristles shorter than primary inner bristles; Ovrlp, degree of overlapping of bristles at the base (0 = not observed: 1 = San overlap; 2 = definite overlap); 
2"9-0, evidence for a secondary outer whorl of short scaly-bristles; 2ne- Oo pata. mie of the secondary outer mites (0 = linear, 1 = narrowly triangular, 2 = 
lanceolate); % = length compared to 1° inner bristles; No. Br= ap Jate number of mid to long bristles (2"¢ inner, 1° outer and inner whorls; small sample 
size); percents listed are bristle lengths compared to the length bathe: primary inner series. 


Taxon Clv Clv- Lgth Ovrlp 2rd. 2"4_9 24.9 No. Comments 
Tap 0 wid % Br 


Heterotheca (x=9) 


sect. Heterotheca (di phic; ray cypselae nearly always lack pappus) 
grandiflora 1 y y ] y 0-1 5-15% 50-60 2°¢ inner bristles few, 40-70% 
H. inuloides 
var. inuloides 1-2 y y 1 y 1 5-15% 60-65 2°? inner bristles many, 35-65% 
var. roseum ] y y 1 y O-1 5-10% 60-65 2° inner bristles few, 50-70% 
var. viridis Q-1 y y ] y 1 5-10% 60-65 2” inner bristles few, 70-80% 
H.leptoglossa 2 y y 1 y 1 5-15% 35-45 2” inner bristles few, 70-80% 


grading into 1° outer bristles 
H. subaxillaris 


subsp. latifolia ] y y 2 y 0-1 5-10% 25-30 2” outer bristles 5-10%; 2™ inner 
bristles O-very few, 60-70% 
subsp. subaxillaris ] y y ] y O-1 5-15% 30-45 2™ outer bristles 5~10%; 2" inner 


bristles few, 40-70% 
sect. Ammodia (rayless) 


H. oregona 
var. compacta ] y y ] y @ 5-8% 60-75 2™ outer bristles obscure; 
2" inner bristles few, 50% 
var. oregona ] y y 2 y 0 5-10% 75-85 2” outer bristles obscure: 


2° inner bristles few, 50% 


SLIVUL Sfidd¥d JVNIGISdOSYAHD ‘J1dW3S 


605 


TABLE 1. cont. 


OLS 


Taxon Clv Clv- Lgth Ovrlp 2nd. 24.9 24.9 No. Comments 
Tap 0 wid % Br 
var. rudis 1 y y | y 0 5-10% 50-55 2°° outer bristles few; 2°? inner 
bristles few, 40-70% 
var. scaberrima 1 y y | y 0 15-30% 60-70 2" inner bristles few, 40-70%, 


grade into 1° outer bristles 
sect. Phyllotheca 
H. brandegei (rayless) 1 y y 1 y 0-1 10-15% 35-55 2™ inner 70-80%, grade into 1° 
outer bristles 


H. canescens 1 y y 1 y 1 5-10% 25-40 2.4 inner bristles few, 50-70% 
H.camporum 
var. camporum 1 y y 1 y 0-1 10-15% 30-45 2" inner bristles O-few, 70% 
var. glandulissimum 2 y y 1 y 0-1 5-10% 30-45 2° inner bristles few, 50-70% 
H. fulcrata 
var.amplifolia O-1 y y 1 y O-1 10-15% 35-45 2™ inner bristles few, 55-75% 
var. arizonica ] y y ] y O-1 10-20% 35-45 2” inner bristles very few, 60-65% 
var. fulcrata 0-1 y y 1 y 1-1+ 10-15% 25-45 2° inner bristles O-few, 70-80% 
var. senilis 1 y y 2 y 1 5-15% 28-45 2°4 inner bristles few, 70-85% 
grade into 1° outer bristles 
H. gypsophila 1-2 y y ] y 0 5-10% 30-40 2" inner bristles few, 70-85%, 
grade into 1° outer bristles 
H. jonesil 1 y y 2 y 1 5-15% 25-35 2” inner bristles O-few, 70% 
H.marginata 0-1 y y 1 y 0-1 10-15% 40-52 24 inner bristles very few, 70-80% 
H. mexicana 0-1 y y 1 y 0-1 10-15% 40-60 2° inner bristles few, 40-60% 
H. monarchensis 1 y y 1 y 1+ 10-15% 25-40 2° inner bristles O-very few, 70% 
H. mucronata 
var. mucronata 1 y y 1 y 0-1 5-10% 35-50 2° inner bristles few, 80-85% 
var. harmsiana ] y y 1 y 0-1 10-40% 35-40 2™ outer bristles can be long; 


2” inner bristles few, 70-80% 


(L)@2 WaIs/9Y¥O' LY 


TABLE 1.cont. 


Taxon 


H. pumila 

H. rutteri 

H. sessiliflora 
subsp. bolanderi 
subsp. echioides 
var. bolanderioides 
var. echioides 


var. camphorata 
subsp. fastigiata 
var. fastigiata 


var. sanjacintensis 


subsp. sessiliflora 
H. shevockil 
H. stenophylla 

var. angustifolia 


var. stenophylla 


H. thiniicola 


Clv Clv- Lgth Ovrlp 2nd 24.9 24.0 No. Comments 

Tap 0 wid % Br 

0) - y 2 y 0 10-20% 35-45 2™ inner bristles very few, 75-80% 

0 = y 1 y 0-1 20-25% 35-46 2™ inner bristles very few, 75-80% 

1 y y 1 y 1 5-15% 50-70 2” inner bristles few-many, 55-85% 

2 y y 1 y 1 X-X% 35-60 2°? inner bristles few, 55-70% 

1 y y 1 y 1 5-15% 25-50 2" inner bristles 0 or few—many, 
60-70%, some fruits have 
significantly few bristles and 
lack 2" inner whorl 

2 y y 1 y 1 5-15% 35-45 2°? inner bristles few, 60-70% 

0-1 y y 2 y 0 5-30% 50-70 2" inner bristles many, 45-85%, 
grade into 2° outer and 1° outer 

whorls 

] y y 2 y 0 5-30% 50-80 2” inner bristles Lo) 45- ica 
grade into 2" outer 
inner whorls 

1-2 y y 2 y 1+ 5-10% 45-50 2™¢ inner bristles few, 50-70% 

1 y y 0) y O-1+ 5-10% 35-45 2™ inner bristles few, 60-65% 

] y y 1 y 0-1 5-15% 35-45 2°? inner bristles few, 50-70%, 
grade into 1° inner bristles 

1 y y 1 y 0) 15-10% 25-42 2°? inner bristles few, 60-70%, 
grade into 1° inner bristles 

1 y y 2 y 0 10-25% 25-35 2°? inner bristles many, 30-60%, 


grade into 2"° outer and 1° inner 


SLIVUL SddWd JVNIGISdOSYAH) ‘J1dWAS 


LLS 


TABLE 1. cont. 


Taxon 


H. villosa 
var. ballardii 
var. depressa 
var. foliosa 
var. minor 
var.nana 
var. pedunculata 
var. scabra 
var. sierrablancensis 


var. villosa 


H. viscida 
H. zionensis 
Noticastrum (x=9) 
acuminatum 
N.calvatum 
N. diffusum 


N. gnaphaloi oides 


N. sericeum 


Clv Clv- Lgth Ovrlp 2nd. 24.9 2nd_Q No. Comments 

Tap 0 wid % Br 

] y y ] y O-1 5-10% 35-45 27 inner bristles few, 60-70%% 

1 y y 1 y 0) 10-15% 5-45 2° inner bristles few, 50-65% 

1-2 y y 0 y 0-1 8-15% 35-45 2” inner bristles few, 60-65% 

] y y 2 y 0-1 5-15% 30-45 2” inner bristles few, 60-70% 

1-2 y y 0) y 0-1 10-15% 28-40 2” inner bristles O-very few, 65% 

2 y y 1 y 1 10-25% 25-45 2°° inner bristles O-very few, 80% 

| y y 0) y 0-1 8-15% 35-45 2°° inner bristles very few, 60-70%, 
grade into 1° inner bristles 

1-2 y y 1 y 0-1 10-15% 35-45 2° inner bristles 70-80%, grade 
into 1° inner bristles 

1 y y 0 y 0) 5-15 30-50 2” inner bristles 60-70%, grade 
into 1° inner bristles 

1-2 y y 0 y 1 5-15% 23-40 2° inner bristles very few, 50-60% 

1 y y 1 y ) 10-15% 30-45 2™ inner bristles few, 50% 

0-1 y y 2 y 0 5-20% 75-90 2™ inner bristles many, 65-75% 

0-1 y y 2 y 0 5-10% 90-100 2” inner bristles few, 80% 

0-1 y y 2 y 0 10-30% 5-80 barbs of bristles tinted red; 2"° 
inner bristles few, 60-80% 

0-1 y y 2 y 0 5-10% 100-115 2” inner bristles few, 80% 

0-1 y y 2 y 0 10-20% 45-50 2™ inner bristles few, 40-80% 

0-1 y y 2 y 0 10-30% 80-90 2” inner bristles few, 80-85% 

1 y y 2 y 0 5-15% 75-80 2" inner bristles few, 70-85% 

0-1 y y 2 y 0 10-20% 120-125 2°? inner bristles few, 70-80% 


ZLS 


(L)@Z VOIS/DYO"LINa 


TABLE 1.cont. 


Taxon (lv Clv- Lgth Ovrlp 2nd. 24.0 24. No. Comments 


Tomentaurum (unknown) 
T.nivea 


0-1 y y 2 y 0 10-20% 75-85 2” inner bristles few to many, 
60-70% 
Pityopsis (x=9) 
sect. Pityopsis 
P falcata 0-1 y y 1 y 0 10-15% 30-40 2° inner bristles O-few, 45-80% 
P flexuosa 1 y y 1 y ) 5-10% 30-50 2°° inner bristles 0O-few, 45-85% 
P pinifolia 1 y y 2 y 1 10-15% 25-30 2” inner bristles not seen 
thii 0-1 y y ] y 1 5-15% 25-35 2™ inner bristles, few, 40-85% 
sect. Graminifoliae 
P. r 
var. adenolepis 0-1 y y 2 O-1 10-20% 25-35 2™ inner bristles O-few, 70-80% 
var. aspera 0-1 y y ] y 0-1 5-15% 30-35 2” inner bristles few, 70-80% 
P. graminifolia 
var. aequilifolia 0 y y 0) y 0-1 10-15% 40-45 2° inner bristles 0O-few, 70-80% 
var. graminifolia 1 y y 2 y 0 10-20% 32-40 2° inner bristles few, 75-85% 
var. latifolia 0 y y 1 y 0 5-15% 35-45 2°¢ inner bristles 0O-few, 60-85% 
var. tenuifolia 0 y y 1 y 0-1 5-15% 35-45 2°° inner bristles O-very few, 70- 
85%, grading into 1° outer bristles 
var. tracyi 0-1 y y 0 y 0-1 5-10% 35-45 2 inner bristles few, 50-85% 
P oligantha 0-1 - y 1 0-1 5-10% 25-40 2” inner bristles O-very few, 
75-85% 
Croptilon (x=7,6,5,4) 
C. hookerianu 
var. graniticum (x=7) 0 = y 0 y 0 5-20% 30-35 2°? whorls absent or very rarely a 


short linear bristle present 


SLIVYL SMdd¥d JVNIGISCOSYAHD ‘31dW3S 


ELS 


Tasle 1. cont. 


Taxon 


var. hookeri (x=6) 


var. validum (x=5) 


C.rigidifolium (x=6, 5) 


C. divaricatum (x=4) 


Bradburia (x=4,3) 
irtella 


B.p 


ilosa 


(lv Clv- Lgth Ovrip ana 24.9 2n4_() No. Comments 

Tap 0 wid % Br 

) ~ y 0 y ) 5-10% 25-30 ° outer vityetes a 2” inner 
oe 5 O-few 

0 7 y 0 y 0 (5-10%) 25-30 2°° outer vac Ae rarely 
vestigial (or abnormal short 1° 
bristle) 

) - y ) n 0) 5-10% 25-30 2” out bristles O-few; 2" outer 
bristles few 35-75%, grade into 
1° bristles 

0 - y 1 n - = 25-30 2” outer bristles not seen; 1° outer 
bristles 70-80%, sometimes grade 

nto 1° in 
O0-1(-2) y y 2 y 2 10-15% 30-36 Ey flo ae inner bristles few, 
0-60%; 1 r 70-95%, grade 

into 1° nae see text for comments 
on pigmentation: disc florets—re- 
duced pappus, 0-5 short scales to 
long broad-based bristles 

O(-1)  y y 0 y 2 8-20% 18-25 2™ outer whorl of large, obvious to 


naked eye, lanceolate-oblon 
scales 0.1-0.2 mm wide; 2°? inner 
bristles O-few, 75-85%, usually 
absent, grade into 1° bristles; 1° 


comments on pigmentation. 


Pls 


(L)@Z VOIS/9YO' LYS 


TABLE 1.cont. 


Taxon 


Clv 


Clv- 


Lgth 


Ovrip 


2nd. 


2n4.9 


24.9 


Comments 


Tap 


0 


wid 


% 


Br 


Chrysopsis (x=5, 4 and x, = 9; most species lack 2" inner bristles and have few primary outer bristles) 
C.de 2 y y y 0 15-25% 


laneyi 
C.floridana 
dfreyi 
var. godfreyi 
var. viridis 
C. gossypina (x, = 9) 
subsp. cru/seana 
subsp. gossypina 
subsp. hyssopitolia 
C. highlandsensis 
C. lanuginosa 
C. latisquamea 
C. linearifolia 
subsp. dressii 
subsp. linearifolia 
C.mariana (x = 4) 


C. scabrella 

C. subulata 
Osbertia (x=5) 

O. bartlettii 


] 


y 


<< 


Me. Oe eS 


y 


<— os 


ain ai ai a a 


] 


—- Oo —- 000 © 


y 


Se Ss Ss OS 


0 


10-15% 


10% 
5-10% 


10-15% 
10-15% 


10-15% 


15-30% 
—15% 


25-30 


30-35 
30-35 


25-35 
20-25 


20-25 


2 inner bristles not seen 
2™ inner bristles not seen 


2” inner bristles not seen 
2” inner bristles not seen 


2° inner bristles not seen 
2° inner bristles not seen 
2° inner bristles not seen 
2™ inner bristles very few, 55-60% 
2™ inner bristles not seen 
2™ inner bristles not seen 


2™ inner bristles not seen 

2™ inner bristles not seen 

2” outer bristles no wider than in 
some other species of genus 
(sample includes 2x, 4x, 6x, 8x); 

2” outer bristles absent; 1° outer 
bristles 0O-few 

2” inner bristles not seen 

2™ inner bristles not seen 


1° vary in length (80-100%); one 
fruit had 1 fine 2°° bristle 25% the 


SLIVHL SNdd¥d JVNIGISdOSYAHD ‘1d WS 


SLs 


T l.cont 
Taxon Clv Clv- Lgth Ovrip 2 24.9 274.9 No. Comments 
Tap 0 wid % Br 
length of the longest 1° bristles 
O. chihuahuana 1-2 y y 0-1 y 0 12% 25-30 2° outer bristles few; 2"° inne 
bristles 45-65% of the length of 
the longest 1° bristles; 1° outer 
bristles 85-95%, grade into 1° 
inner bristles 
O. stolonifera 0-1 - y 1 y 0 5-40% 25-30 2™ outer bristles grade into 2 


inner bristles; 2" inner bristles 50— 
70% of the length of the longest 1° 
bristles; 1° outer bristles 90-95% 
grade into weakly clavate 1° inner 
bristles 


(L)@2 VaIs/9¥O' LIYE 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 517 


inner bristles; primary inner bristles very weakly (Fig. 1B) to moderately clav- 
ate (Fig. LK), 3.4-10 mm long. 

Noticastrum (x = 9; eight of 19 species examined).—Cypselae straw col- 
ored to brown, fusiform, 4-6 mm long, sparsely to moderately strigose, some- 
times stipitate glandular distally (Fig. 2B-C) or over entire surface (Fig. 2H); 
ribs, 16-22, surface between ribs usually golden-brown, translucent, resin filled; 
pappi quadruple, each series usually grading into next, 45-125 mid to long 
bristles, averaging about 87 per fruit for the genus, straw colored, orange-rust 
or purplish (e.g., N. diffusum; Fig. 2A), yellow-rust pigment in shaft over entire 
length, the red pigment concentrated in the barb tips (Fig. 2F), barbs evenly 
arranged around the round shaft along the entire length of the bristles; sec- 
ondary outer series obvious to inconspicuous, linear triangular scales (Figs. 2N) 
to linear bristles (Fig. 2B), usually 5-20(-30)% the length of the longest pri- 
mary bristles; secondary inner bristles many, 40-85% the length of the longest 
primary bristles (Fig. 2E); primary inner bristles many, 85-95% the length of 
the primary inner bristles; primary inner bristles very weakly (Fig. 2D) to some- 
times moderately clavate, 7-14 mm long, much exceeding the disc corolla lobes. 

Tomentaurum (x = unknown; monotypic, IT. nivea examined).—Cypscelae 
fusiform, 4-5 mm long, densely strigose (Figs. 3B-C); ribs 10-18, shallow, ob- 
scured by hairs; pappi quadruple, each series usually grading into next, 75-85 
mid to long bristles, averaging about 80 per fruit, straw colored, barbs evenly 
arranged around the round shaft along the entire length of the bristles; sec- 
ondary outer series obvious to inconspicuous, very linear triangular scales to 
tapering bristles (Figs. 3H-D, 10-20% the length of the primary inner bristles; 
secondary inner bristles few to many, 40-85% the length of the longest pri- 
mary bristles (Fig. 3G); primary inner bristles many, 85-95% the length of the 
primary inner bristles (Fig. 3D); primary inner bristles very weakly to weakly 
clavate (Fig. 3E-F), 10-11 mm long. 

Pityopsis (x = 9; all seven species, all 11 taxa examined).—Cypselae straw 
colored to often dark reddish-brown, narrowly to broadly fusiform, 3-4 mm 
long, ribs 8-10, sparsely to moderately densely strigose; pappi quadruple or 
sometimes triple through loss of the secondary inner series, 25-46 mid to long 
bristles, averaging about 37 per fruit for the genus, light tan to straw colored, 
sometime pale rust colored proximally, barbs evenly arranged around the round 
shaft along the entire length of the bristles; secondary outer series of linear 
(Figs. +E, G arrows) to broad (Figs. 4+K-L), barbed scales, 5-15(-20)% the length 
of the primary inner bristles; secondary inner series of tapering bristles, few, 
45-85% the length of primary inner bristles; primary outer series of tapering 
bristles 90-95% of primary inner bristles (Figs. 4C-D, E H; arrows); primary 
inner series of weakly clavate bristles (Fig. 4D), 4-9 mm long. 

Croptilon (x = 7, 6, 5, 4; all three species, all five taxa examined).—Cy pselae 
straw colored to brown or reddish-brown, sometimes mottled, fusiform to nar- 


518 BRIT.ORG/SIDA 22(1) 


Fic. 2 papy i ticast : 

(Zardini 1094 TEX). A. Fruit with rusty reddish antl 
secondary outer bristles (arrow) and the bases longer seri istl leb mm. €. SEM of stipitate glands 
(arrows) on upper fruit body; scale il = 100 um. D. lips of primary inner series srs, E. ee Ui seeunealy inner 


t ? Peet 


(H hbach 52879 MO). H. Fruit t | I istle ( ).1 Detail of multi- 
ribbed ues mony with stipttate ganas kart): - N. macrocephalum (Montes 14942 MO). I. Maule fruit are Im- 
f K.T clavate to very v p of primary 
r Pa 
tert tl 1 PP fal y rina tle f 1 y bristle and Uppel mid portions 


(open arrow). 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 519 


Fic. 2.5 . 1 Pema re £T. ° 1° Leisite (Dre, ln TIZAL RAN) lah Tmmin A-B =0.5 mm 


in H. A. Nearly fruit with lary outer series (1°o), secondary inner series (2°i), primary outer series (1°0) 
J id . id (1 O°V bL 7-4] . J* 7 J R Eruit hndv C€ AA A 4 £f, Hy | J nm TT: £| - 1 car fl | 
r é co r é f vf 

a few are broken off. E-F. Tips of primary inner bristles. G. Tip of dary i bristle ( ) and mid porti f 


four primary bristles. H-I. Secondary outer series bristles. 


rowly obconic, 2-3.2 mm long, ribs 10-20, sometimes either the ribs or faces 
between them golden-brown to brown translucent, resin filled, sparsely to 
moderately densely strigose (Figs. 4A, G, K); pappi quadruple, triple, double, or 
sometimes single, 25-35 mid to long bristles, averaging about 28 per fruit for 
the genus, somewhat to dark rust colored in shaft, darkest proximally, becom- 
ing paler to whitish distally, barbs not pigmented, barbs evenly arranged around 
the round shaft along the entire length of the bristles; secondary outer series of 
linear bristle-like scales, O-few (Figs. 5D-E, I-J, N-O), 5-10(-20)% the length of 
the primary inner bristles (Figs. 5D-F, I-J, O); secondary inner series of taper- 
ing bristles, absent or rarely 1-2, 50-70% the length of primary inner bristles 
(Fig. 5A, M; arrow) or these just atypically short primary outer series bristles; 
primary outer series of tapering bristles 80-95% of primary inner bristles, grad- 
ing into primary inner bristles (Figs. 5A, 5G, 5K-L); primary inner whorl of non- 
clavate/very weakly clavate (Fig. 5B) to weakly clavate bristles (Fig. 5L), 4-9 
mim long. 

Croptilon has been treated as Haplopappus sect. Isopappus (Hall 1928; Smith 
1965) and described as having a pappus with “a single series of equal (or nearly 


520 BRIT.ORG/SIDA 22(1) 


Fic. 4. Frui its of Pityopsis, disc frui let 1mm in A-B and I,= 100 Lim inJ, = 0.5 mm in K, and 


= 0.1 mmin L.A. P graminifolia var. - aequilifolia, mature fruit (Semple et al. 3992 WAT). B—E. P. falcata (Semple 3366 
WAT). B. au aay: C- 2 Tips of weakly caval primary inner bristles and attenuate, shorter primary outer bristles 
(arrows).E y | Es sels ead a re F. Tips of primary inner and 
outer bristles G. Bri ly secondary outer i | 0537 WAT, tips of primary inner 
and outer (arrows) series bristles. \- L. ane var. mdenelepD (empleet a wor, lL. Maullte murchad ys = oe NS 
J. Clavate tips of primary inner series bristles. y highligl 

K) and proximal portions primary series ae 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 


Fic. 5. Fruit and pappus traits of Croptifon, disc fruits; scale bars = 1 mm. A-D. C. hookerianum var. hookerianum. A 
Mature fruit (Semple & Brouillet 3363 WAT).B—D. Turner et al. 46 TEX-LL.B. Tips of primary bristles. C. Tips of secondary 
inner bristle and upper mid portions of two longer Hee D. Tip of suis outer arisile, E.C, negkeanum var. 
graniticum, secondary outer bristle (E.B. Smith 622 TEX). F. 

4259 TEX). G—J. C. rigidifolium (G and |, Cory 11886 TEX; ee Nesom 5216 TEX, SEMs). G. Mature fruit. H. tine of pri- 
mary bristles. I-J. Bases of primary bristles and secondary outer buses fartaws). K- iM C iver Cemnle 


Cimiclowsse0n) WAT). K. Mature uth a iy of panany bristles. M 


> 


522 BRIT.ORG/SIDA 22(1) 


equal) capillary bristles, more or less ferruginous” (Smith 1965). Nesom (2000) 
as noted in the introduction also described the pappus as being ina single se- 
ries. The results presented here indicate that the pappus neither consists of a 
single series nor are the bristles equal or nearly equal. The higher base number 
taxa (e.g., C. hookerianum var. graniticum with x=7 ) are more likely to have 
three or four series of bristles, while the lowest base number taxon, C. 
divaricatum with x = 4, may have only 1-2 series. The gradation in lengths of 
longer bristles and the lack of clearly clavate bristle tips results in uncertainty 
in determining the number of series present. The conclusion that multiple se- 
ries are present is based on observations of many fruits, some of which have 
longer bristles of 2-3 distinct lengths. The much shorter outer bristles clearly 
exterior to the long bristles are essentially the same as the secondary outer se- 
ries scales/bristles of some species of genera long recognized to have a short 
outer series, e.g. Heterotheca, Noticastrum, Pityopsis and Chrysopsis. The small 
fruit size generally correlates with a lower number of bristles and very few 
bristles in some series. 

Bradburia (x = 4, 3; two species with very different pappi traits).—1) B. 
hirtella: ray florets fertile, disc florets functionally staminate, ovary aborts; ray 
floret cypselae straw colored to brown or reddish-brown, obconic, + triangular 
in cross-section, ca. 2mm long, 5-10 shallow ribs per face, moderately strigose, 
more densely so along angles (Fig. 6A); pappi quadruple or triple, the series grad- 
ing into each other, 18-26 mid to long, long-barbellate bristles (Fig. 6A), densely 
pigmented red proximally to only the barbs pigmented one third to one half 
the length, pale yellow to pale rust grading to white distally; secondary outer 
bristles linear, 5-20% the length of the longest inner bristles (Fig. 6B), barbs 
evenly arranged around the round shaft along the entire length of the bristles, 
uneven on the distal portion of the longer bristles; secondary inner bristles grad- 
ing from secondary outer into primary outer, tapering, 25-85% the length of 
primary inner bristles; primary inner bristles round proximally, flattened dis- 
tally and weakly to moderately clavate (Fig. 6C), 2.5-3.5 mm; mid to longer 
bristles pigmented proximally, the red pigment concentrated in the barbs (Figs. 
OB, D). Disc floret pappi reduced to 0-5 scales and/or broad-based proximally 
flattened bristles (Figs. 0G-J), the longest weakly clavate (Fig. 6H). — 2) B. pilosa: 
ray and disc pappi similar, cypselae obconic, slightly compressed, sparsely stri- 
gose, 10-20 shallow ribs; pappi triple or double (Fig. 7K); secondary outer scales 
linear to lanceolate or oblong (Fig. 6L), 0.05-0.2 mm wide, barbellate proximally, 
margins jagged, especially distally; secondary outer bristles absent, primary 
outer bristles tapering, 0-2, 75-85% the length of the inner bristles; primary 
inner bristles tapering or only very remotely clavate; all longer bristles pig- 
mented proximally with the concentration decreasing distally, yellow to rust, 
pigment in shaft, not barbs. 

The combination of broad, whitish, light-reflecting scales and proximally 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 523 


Saganaeier* 4 
of 
- : 


Fic. 6. Fruit and papy tae ds Bigapuna, scale bars = ao mm. oe J. B. hirtella (Nesom 7470 WAT). A-E. Ray fruit. B. 
g b g g outer brist tle caivow); e Tip of Ae! fattened Hong pet 

L 4 1 D MAtd Ld at ¥ 1 aL 4; TL L iT 
devel J. Functionally inate disc floret. F. Floret with Jary le ( ") 1 two long 
broad-b ad hrictles G.B f f] ith | j | H. Tit ra he GA. Tiy oe 


in G. J. Details of scales bases. K-L. B. pilosa, disc fruit (Semple & Brammall 2753 WAT). K. Mature fruit. L. Secondary 
outer series scales. 


524 BRIT.ORG/SIDA 22(1) 


darker-pigmented, long bristles as background makes the secondary outer se- 
ries of Brad buria pilosa obvious to the naked eye. The narrower to mid-width 
scales are similar to broader scales in some species of Chrysopsis and 
Heterotheca. The widths of the secondary outer scales/bristles in Heterotheca 
and Chrysopsis form a continuum that overlaps at the wide end with the nar- 
rower width scales of B. pilosa. The broader scales of B. pilosa are unique in the 
subtribe. The broad scaly bases of the few, long bristles attached to the aborted 
ovary of the disc florets of B. hirtella are also unique in the subtribe. Possibly, 
the genes controlling normal scale and bristle development have been expressed 
together in single structures rather than separately in different series of struc- 
tures in disc floret pappus development. Therefore, but in different ways, the 
two species of Bradburia have broader pappi members than found in all other 
genera in the subtribe. 

The phylogenetic position of Bradburia is uncertain. Nesom (199la, d) in- 
cluded the two species of Brad buriain Chrysopsis sect. Brad buria. Semple (1996) 
transferred C. pilosa to Bradburia as defined here. In the cladistic study by 
Semple and Tebby (1999), Bradburia grouped with Croptilon and Heterotheca, 
while Chrysopsis grouped with Pityopsis, Noticastrum, and Tomentaurum. 
Based on pappus traits, Bradburia is no more similar to Croptilon than to 
Chrysopsis. Pigmentation of bristles is similar in Bradburia and Croptilon, but 
a well developed secondary outer series is shared by Brad buria and Chrysopsis, 
although the scales are much broader on average in B. pilosa than in any spe- 
cies of Chrysopsis. A DNA sequence based phylogeny of the subtribe has not 
been published. Until such a study is completed, the phylogenetic position of 
Bradburia within the subtribe remains uncertain, although it is likely derived 
from either the Croptilon or Chrysopsis lineages, unless a separate dysploid se- 
ries from x = 9 is hypothesized. 

Chrysopsis (x = 5, 4, x2 = 9; all taxa examined, 11 species, 15 taxa).— 
Cypselae straw colored or rarely dark red-purple, compressed obconic, smooth 
(Fig. 7G) or with 1-10 shallow ribs and sometimes 1-5, yellow to red brown, 
clavate, translucent ridges per side (Fig. 7D), sparsely to densely strigose, 1.5-3 
mm, pappi triple or rarely quadruple, sometimes double in C. mariana, barbs 
evenly arranged around the round shaft along the entire length of the bristles; 
secondary outer series of linear (Fig. 7F) to narrowly triangular, jagged-edged 
scales (Figs. 7C, 7L), 0.4-1.4 mm, 5-15(25)% the length of the primary inner 
bristles, 20-40 mid to long barbellate bristles in 1-2(-3) series, whitish to straw 
colored; secondary inner series usually absent, rarely 1-3 tapering bristles 45- 
85% the length of primary inner bristles; primary outer series of tapering 
bristles 85-95% of primary inner bristles (Figs. 7B, E, larrow, K); primary inner 
series of weakly to moderately clavate bristles (Figs. 7E, I-J), 4-7 mm long. 

Some fruits of C. mariana (x = 4) have a true double pappus of just short 
secondary outer series scales and long, clavate primary inner series bristles (Fig. 


— 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 525 


Fic. 7. Frui J its of Ct is, disc frui let Sl 0.5 mm in L.A-C. C. latisquamea 
BW aah 2753 WAT) A. Mature fruit B. Tips of bri C.5 y De ms scales D. Fruit epady 
jish-brown resin- filled rib Co (Semple et al. 312 pap 
pus, C. delanevi (Semple & Semple 7476 WAT).F. Li bristle-li le ( ) ifolia (Godfi 
75759 WAT). . at ioe une Cis godireyi (Semple & Godfrey 3149 wal): ae C. mariana Cemple & 
Chmielewski 6214 WA I uter series; 


0—2 primary outer bristles on other fruits SOU l. ips of primary inner bristles and one primary outer bristle 


(arrow). J. Linear t narrowly trian gular y 


526 BRIT.ORG/SIDA 22(1) 


7H, 7L); others fruits produced by the same individual have a few tapering pri- 
mary outer bristles (Fig. 71. This is true for fruits at all four ploidy levels 2n=8, 
16, 24, 32: Semple & Chinnappa 1986). In his cladistic study, Nesom (1991a) 
scored the secondary outer scales of both Bradburia pilosa and C. mariana as 
“(1) scales”, but the latter has scales that are very narrowly triangular to only 
narrowly triangular on the same fruit like some other species of Chrysopsis, 
not like those of B. pilosa. 

Osbertia (x = 5; three o 
to brown, cylindrical-fusiform, 1.0-1.5 mm long, sparsely strigose, ribs 8-16, su- 
perticial (Figs. 8N, T), sometimes either the ribs or faces between them golden- 
brown to brown translucent; pappi quadruple, triple, double (Fig. 8M) or single 
and rarely vestigially triple, the series distinct or grading into each other (Fig. 
8A), 20-35 bristles, barbs evenly arranged around the round shaft along the 
entire length of the bristles, barbs sometimes small and less obvious; second- 
ary outer bristles linear, 3-10, sometimes grading into secondary inner bristles, 
absent in O. bartlettii or very rarely one linear, fine, short bristle (Figs. 8R-S), 5- 
40% the length of the inner long bristles; secondary inner bristles tapering, 
fine, linear, 5-10, 45-70% the length of primary inner bristles, absent in O. 
bartlettii; primary outer bristles fewer than and often grading into the primary 
inner bristles, 85-95% the length of the primary inner; primary inner bristles 
weakly tosometimes moderately clavate tapering (Figs. 8C, I-]), or in O. bartletii 
tapering and subequal or the shortest 80% if all long bristles are included ina 
single series, 2.5-7.5 mm long. 

The pappus of Osbertia differs in the three species, but in general all have 
rather linear and short-barbed bristles. The four series are most distinct length- 
wise in O. chihuahuana (Figs. H-L) and least distinct or reduced to two or one 
series in O. bartlettii. In the type species C. stolonifera, the secondary outer and 
inner series grade together, as do the primary outer and inner series, but the 
secondary and primary series are clearly of different lengths (Figs. A-B). The 
two secondary outer series could be interpreted as a single series highly vari- 
able in length, and the primary outer and inner could be interpreted asa single 
series grading from non-clavate shorter bristles to weakly clavate longer bristles. 
Overall there is a spiral arrangement from very short, fine, outer bristles to long 
coarser inner bristles, with the outer members of the series clearly placed ex- 
ternally to and overlapping the longer inner members of the series. 

Previous authors have considered the pappus of Osbertia to be simpler than 
described above. Turner and Sundberg (1986) described the pappus of Osbertia 
as having “numerous setae ina single series” 4+-6 mm long. Nesom (2000) noted 
that O. chihuahuana Turner & Sundberg had “a short, outer series commonly 
present” (i.e.,a double pappus) but did not describe the individual bristles or 
scales. Nesom (1991c) stated that pappus of O. bartlettii was “uniseriate, of 15- 
20 barbellate bristles without an outer series.” The single short outer bristle 


Ss 


three species examined) — Cypselae straw-colored 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 527 


Fic. 8. Frui i Osbertia, disc frui le bar= 1mm. A-G. 0. stolonifera (Ton 1027 TEX-LL). A. Imma- 


ture fruit. B. Distal portion oflonder iste. hs au A) ELLA vowen inner bristles. D-E. Tips of mid length 
econdary yl a G.S pectively. H— te Me chihu ag 


7 


sare edo 2232 TEX Distal half of ( indi possibl ).1 tal portion of 
primary bristles. J. a tay ier K. Tip of possibl lary bristles (center in s L. Linear sec- 
ondary outer bristle (arrow). M—T. 296 WAT; SEM images, Muller 2854 UC.M. Immature fruit with 


disc corolla attached. N. Fruit body sam oe Q. Tips of long primary bristles. R-S.Tip and base of rarely seen short outer 


bristle, respective 


528 BRIT.ORG/SIDA 22(1 


= 


reported above was found on one fruit of the few present on the single collec- 
tion of the species at WAT (Nesom 6296), but none were visible in the material 
from UC used for the SEM examination (Fig. 8H). 

The phylogenetic position of Osbertia is uncertain and the genus only ten- 
tatively has been included here in the Chrysopsidinae. It was included in 
Haplopappus sensu lato by Hall (1928), which sensu stricto is part of the 
Machaerantherinae Nesom. Nesom (1991c) placed Osbertia in the 
Chrysopsidinae, and Nesom (2000) placed it close to Chrysopsis. In the RFLP 
DNA study by Lane et al. (1996), Osbertia stolonifera was placed basal to the 
clade including Ericameria/Macronema, Xylothamia, and Tonestus but was not 
included in the ITS DNA sequence study by Beck et al. (2004). In the latter study, 
Ericameria, Xylothamia, and Tonestus were each placed in different clades sepa- 
rate from the clade including Heterotheca villosa and Chrysopsis gossypina. The 
phylogenetic position of Osbertia within the North American clade needs ad- 
ditional study. On the basis of pappus traits, Osbertia fits into the subtribe 
Chrysopsidinae as a derived lower base number taxon with a reduced number 
of bristles, sometimes reduced number of series, and derived non-clavate to 
weakly clavate longer bristles. Further study is needed to determine if such 
traits would exclude it from other proposed subtribes. 


SYSTEMATICS IMPLICATIONS 


There is considerable variation in the pappi traits of taxa within the 
Chrysopsidinae, which can not be defined in part on the basis of usually hav- 
ing a double pappus. Even within individual genera there is variation in the 
number of series, how distinct the series are from each other, and the number 
of long bristles per fruit. There are patterns to the variation such that a “norm” 
for each genus can be given, but these are not diagnostic due to with-genus varia- 
tion and overlapping of ranges of variation between genera. For example, the 
number of bristles in Noticastrum, with a quadruple pappus, is higher com- 
pared to other genera; the bristles in all series are linear; and the tips of the 
longest secondary inner bristles differ little from those of the shorter bristles 
in the primary outer series and secondary inner series. In Heterotheca with a 
usually quadruple pappus, the number of bristles averages less than 
Noticastrum but more than occur on the fruits of the lower base number taxa; 
and there are usually only a few mid-length bristles making up the secondary 
outer whorl. However, in H. oregona the number of bristles is within the range 
found in Noticastrum, but the series are generally discontinuous in length and 
thus do not grade into each other. In other species of Heterotheca, the bristle 
series grade into each other, but there are fewer bristles, e.g., H. thiniicola. In 
Chrysopsis with a usually triple pappus, the number of bristles and the absence 
of secondary inner bristles is fairly consistent; the primary inner bristles usu- 
ally are more obviously clavate than in most species in other genera of the 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 529 


subtribe; and fruits with relatively broad secondary outer scales have some lin- 
ear scales. Chrysopsis mariana with x = 4 has a double pappus or “slightly” triple 
pappus (1-2 primary inner bristles), while the secondary outer scales are like 
other species in the genus and the primary inner bristles are obviously clavate 
(especially when examined at 50-100H). Although pappus descriptions of the 
genera of the Chrysopsidinae need to be longer to encompass the details and 
range of variation in the series of bristles, the individual genera themselves can 
not be diagnosed on the basis of pappus traits alone. Therefore, the details of 
pappus features presented here are likely to be more significant for cladistic/ 
phylogenetic studies then for identifying specimens. The genera with the higher 
base numbers have more pappus elements in more series than do the lower base 
number genera with more derived and simpler pappi. Lastly, when the pappi 
of members of other subtribes have been examined in detail, general trends in 
differences of derived features may emerge. At this point, however, more data 
are needed. 


ACKNOWLEDGMENTS 


This research was supported by a Natural Sciences and Engineering Council of 
Canada Discovery Grant to J.C.S. A preliminary analysis of pappi traits was 
included in a cladistic study of the subtribe done by Laura Woodworth as part 
of a BIOL499 senior research project at the University of. Waterloo. Dwayne 
Estes (TENN) kindly provided data on the pappus of Pityopsis ruthii. The tech- 
nical assistance of Jennifer Hood and Naomi Steenhof is gratefully acknowl- 
edged. 


REFERENCES 


Beck, J.B.,G. Nesom, PJ. Catie, R.K. SMALL, and E.S. ScHILLING. 2004. Is Solidagininae monophyl- 
etic? Taxon 53:691-698. 

Cronaquist, A. 1955.Vascular plants of the Pacific Northwest. Part 5: Compositae. University 
of Washington Press. Seattle. 

Cronaquist, A. 1968. Aster, Chrysopsis, and Heterotheca. In: H.A. Gleason, ed. The new Britton 
and Brown illustrated flora of the northeastern United States and adjacent Canada. 
Hafner Pub. Co., New York. 

Cronauist, A. 1980.Vascular flora of the southeastern United States — Vol. 1 Asteraceae. The 
University of North Carolina Press, Chapel Hill. 

FerNato, M.L. 1950. Gray’s manual of botany 8" Ed. D.Van Nostrand Company, New York. 

Gray, A. 1884. Synoptical flora of North America. Vol. 1, part 2. vison, Blakemean, Taylor & 
Co., New York. 

Hatt, H.M. 1928. The genus fe da a phylogenetic study in the Compositae. Publ. 
Carnegie Inst. Wash. 389:1 

Hoimaren, PK., N.H. Houmeren, and eo BARNETT. 1990. Index herbariorum. Part |. The herbaria 
of the world, ed. 8. Regnum Veg. 120:1-693. 


530 BRIT.ORG/SIDA 22(1) 


Hoop, J.L.A.and J.C. Sempte. 2003. Pappus variation in Solidago (Asteraceae: Astereae). Sida 
20:1617-1630. 

Lane, M.A.,D.R. Morcan, Y. Sux, B.B. Simpson, and R.K. Jansen. 1996. Relationships of North Ameri- 
can genera of Astereae, based on chloroplast DNA restriction site data. /n DJ.N. Hind 
(Editor-in-Chief). Proceeding of the International Compositae Conference, Kew, 1994. Vol. 
1. Systematics (vol.eds DJ.N. Hind and H. Beentje). Royal Botanic Gardens, Kew. 

Nesom, G.L.1991a. Union of Bradburia with Chrysopsis (Asteraceae: Astereae), with a phy- 
logenetic hypothesis for Chrysopsis. Phytologia 71:109-121. 

Nesom, G.L. 1991b.Tomentaurum (Asteraceae: Astereae), a new genus of goldenaster from 
Chihuahua, México. Phytologia 71:128-1 

Nesom, G.L. 1991c. Transfer of Heterotheca bartlettii to Osbertia (Asteraceae: Astereae). 
Phytologia 71:132-135. 

Nesom, G.L.1991d.A phylogenetic hypothesis for the goldenasters (Asteraceae: Astereae). 
Phytologia 71:136-151 

Nesom, G.L. 1994a. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 
193-274 

Nesom, G.L. 1994b. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), 
emphasizing the New World species. Phytologia 77:141-297. 

Nesom, G.L.2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America, 
Central America, the Antilles and Hawaii. Sida, Bot. Misc. No. 20. Botanical Research 
Institute of Texas, Ft.Worth. 

Noyes, R. D. and LH. Rieseperc. 1999. ITS sequence data support a single origin for North 
American Astereae (Asteraceae) and reflect deep geographic divisions in Aster S.L. 
Amer. J. Bot. 86:398-41 2. 

Semete, J.C. 1981. A revision of the goldenaster genus Chrysopsis (Nutt.) Ell. nom. cons. 
(Compositae-Astereae). Rhodora 83:323-384. 

Sempte, J.C. and F.D. Bowers. 1985. A revision of the goldenaster genus Pityopsis Nutt. 
(Compositae: Astereae). Univ. Waterloo Biol. Ser. 29:1—34. 

es J.C.and C.C. CHINNaPPA. 1986. The cytogeography of Chrysopsis mariana (Compositae: 

reae): Survey over the range of the species. Rhodora 88:261-266. 

Sani J.C. and J.L.A. Hoop. 2005. Pappus variation in North American Asters. |. Double, 
triple and quadruple pappus in Symphyotrichum and related aster genera (Asteraceae: 
Astereae). Sida 21:2141-2159. 

Sempte, J.C. and L. Teesy. 1999. A cladistic analysis of subtribe Chrysopsidinae (Asteraceae: 
Astereae). XVI International Botanical Congress, August 1-7, 1999. St. Louis, MO. Ab- 
Stract No. 2852; Poster No. 401. 

Semete, J.C., C. Leeper, C. Leury, and L. Gray. 1988. Heterotheca sect. Ammodia (Compositae: 
Astereae): a multivariate study of H.oregona and specimens of Brewer's (golden)aster. 
Syst. Bot. 13:547-558 

Sempte, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: 
Astereae): the prairie and montane goldenasters of North America. Univ.Waterloo Biol. 
Ser. No. 37:1-164 


SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 


Semete, J.C., 5.B. Hearo, and L. Brouitet. 2002. Cultivated and native asters of Ontario 
Compositae: Astereae):Aster L. (including Asteromoea Blume, Diplactis Raf.and Kalimeris 
(Cass.) Cass.), Callistephus Cass., Galatella Cass., Doellingeria Nees, Oclemena E.L.Greene, 
Eurybia (Cass.) S.F. Gray, Canadanthus Nesom, and Symphyotrichum Nees 
Virgulus Raf.). Univ.Waterloo Biol. Ser.41:1-134. 


Smith, E.B. 1965. Taxonomy of Haplopappus section Isopappus (Compositae). Rhodora 
67:21 7-238. 


— 


=alt 


—— 


— 


including 


Turner, B.L. and S. Sunosera. 1986. Systematic study of Osbertia (Asteraceae-Astereae). PI. 
Syst. Evol. 151:229-239. 

ZAROINI, EM. 1985. Revision del genero Noticastrum (Compositae-Astereae). Rev. Mus. La 
Plata, n.s. 13.83:313-424. 


532 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Bossy J. Warp. 2004. The Plant Hunter’s Garden. (ISBN 0-88192-696-5, hbk.). Timber 
Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. 
(Orders: www.timberpress.com, mail@tim| a he com, 503-227-2878, l- 
800-327-5680, 503-227-3070 fax). $39.95, 340 pp, b&w drawings, color photos, 
bibliography, index, 7 3/8" x 10 3/8". 

If you get excited at the sight of the season's first seed and plant catalogs, you should pick up a copy 


of the new book The Plant Hunters Cares n. Author popey Ward includes the profiles of 32 modern 


} 


day plant hunters many of the plant offerings in those catalogs. The people 


included are from finotighout hie oat and are associated with an assortment of nurseries, horti- 


cultural research associations, and/or small companies offering plants or seeds. The focus of the in- 
cluded plant collectors is very ae native plants, regional plants, ‘ aa plants, rock and alpine 


gardens, plants/seeds from inaccessible locales, unusual or hard to find species or varieties, and even 


collectors with very specific interests, such as bulbs, shade plants, and single genera. 
Each chapter focuses ne plant hunter/company. First off, the person or company is pro- 
filed g nee some oe ios mation on the collector's history and plant interests. Then some 


inl non travels and exploration for new plants is discussed. A conclusion for each includes a 
picture of the collector and company contact information. The bulk of each chapter concentrates on 
the plants collected by, introduced by, grown by and promoted by the various plant hunters. The 
chapters are filled with colorful photos of oa as well as some origin descriptions and growing 


zone boy atien for various plants. Throughout the chapters are brief stories about how some plants 
] 


how the person met someone who introduced or passed along the plant to them. 


The descriptions of the discovery of these plants were a little less adventurous than this reader ex- 
pected, but these discoveries are still interesting to read about. An example of sucha discovery is that 
of the small, white flowered, bronze leaved plant Jabrosa volkmannii by John Watson and Anita Flores, 


found growing in pumice on the side a volcano. There is also interesting information on how these 
hunters cultivate their finds. For instance the book ue a few “rock gardeners” growing species 
in various rock crevices including Panayoti Kelaidis of the Denver Botanical Gardens who designed 
and manages a 450,000 kg, 0.4 ha rock garden with 4000+ plant species. The book concludes with a 
chapter on invasive plants that gives an honest look at w 


> 
a] 


at weeds and invasive species are and the 
variety of vectors of ial AnEFOGEC ion. 


If or | li ho thrill i icipati f new species and vari- 
eties of plants, on will thoroughly enjoy reading this Geek and will benefit [om aie contact cs 
mation supplied for each collector. Travel with these plant I 
naeult 


of discovery, excitement of their plant fin vithin the pages of Ward’s 
The Plant Hunter’s Garden.—Lee eon Herbarium, figvantcat Re nee Institute of Texas, 509 


Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(1): 532. 2006 


NOTES ON TYPES IN APOCYNACEA EAASCEEPIADOIDEAE 
IN CUBAN HERBARIA AND FOUR LECTOTYPIFICATIONS 
IN WEST INDIAN GONOLOBINAE 


Alexander Krings Paul R. Fantz 
sie elt Department of Botany Department of Horticultural Science 
orth Carolina State University North Carolina State University 
Ralei a North Carolina 27695-7612, U.S.A. Raleigh, North Carolina 27695-7609, U.S.A. 
Alexander_Krings@ncsu.edu Paul_Fantz@ncsu.edu 
ABSTRACT 
Four asclepiadaceous types not listed in a previ f the collections of the Instituto 


de Egologia) vy SIStetnanA, Elabaiia< Cuba (H AC) are prov rided, alone witha giscussion of typuication 
t I 


1 Charles V 1] 


[ 
for Gonolobus stephanotrichus Griseb., Orthosia acuminata Griseb., Orthosia oblongata coy and 
b 


Poicilla ovatifolia Grise 


RESUMEN 
a, t ] A ] ] 4 ] ] =| : -j : ae | ] 
oO C L 
de igen de Feit Sse a, Habana, Cuba (HAC), junt di ion de] bl 
tipificacion asociados con las Colecciones Charles Wright. 7 eapouan lectotipificaciones de 
ibn stephanotrichus ee 2 iseb,, y Poicilla 


ovatifolia Griseb. 


A revision in progress by Krings of subtribe Gonolobinae (Apocynaceae: 
Asclepiadoideae) in the West Indies has led to the discovery of four additional 
types at the Instituto de Ecologia y Sistematica, Habana, Cuba (HAC). These 
specimens were not listed ina previous catalogue of types at HAC (Cardenas & 
Herrera 1991). Types in subtribe Gonolobinae were not found at the Jardin 
Botanico Nacional, Habana (HAJB; for a complete list see Gutiérrez et al. 1997). 
The expanded list of Apocynaceae: Asclepiadoideae types at HAC is presented 
in Table 1. 

The asclepiadaceous original material at HAC belongs to two groups: (1) 
species with unproblematic typification due to designations by the original 
author(s) and (2) species requiring lectotypification due to complications in 
collecting practice and lack of holotype designations. 

Typification is unproblematic for Marsdenia micrantha Alain, Matelea 
alainii Woodson, and Marsdenia bicolor Britton & P. Wilson, as these species 
were described relatively recently and as each original author designated holo- 
types. Thus, corresponding material at HAC (Table 1), was identified easily as 
either holotype, isotype, or paratype based on the respective protologues. An 


SIDA 22(1): 533 — 537. 2006 


= 


534 BRIT.ORG/SIDA 22(1) 


Taet E 1.List of Apocynaceae: Asclepiadoideae types at the Instituto de Ecologia y Sistematica, Habana, 
Cuba (HAC). Previously unlisted types (Cardenas & Herrera 1991) are marked by an asterisk. 


(1) Species with unproblematic typification 
“Marsdenia bicolor Britton & P. Wilson, Bulletin of the Tor bok Botanical Club 50:47. 1923. 
Type: CUBA: Léon 10787 (isotype: not seen); Léon 10788 (paral 


SS 


Marsdenia micrantha Alain, Revista de la Sociedad Cubana de Botanica 13:59. 1956, 
Type: CUBA: Al ain, Acuna & Lopez 5678 (HOLOTYPE; IsoTyPE: NOt seen) 


a aia inii San ee Contribuciones Ocasionales del Museo de Historia Natural del Colegio 
Salle’ 4.1956 
ener 5029 (8 TYPE!) 


(2) Species lectotypified herein (see text) 
sae stephanotrichus Griseb., Catalogus plantarum cubensium 177.1866. 
E: CUBA: Wright 2969 (syntyee!) 
*Orthosia acuminata Griseb., Catalogus plantarum cubensium 175, 1866. 
Tyee: CUBA: an (ISOLECTOTYPE!) 
hs ‘thosia ot Griseb., Catalogus plantarum cubensium 176. 1866. 
Type: CUBA: Wi ight 2967 (sc LECTOTYPE!) 


*Poicilla ovatifolia Griseb., Catalogus plantarum cubensium 177.1866. 
Type: CUBA: Wright 2965 (syntyee!) 


isotype of Marsdenia bicolor was reported previously by Cardenas and Her- 
rera (1991), but not seen as part of this study. A paratype (Léon 10788) at HAC 
was not listed by Cardenas and Herrera (1991). 

Grisebach (1866) described a number of taxa based on material collected 
by Charles Wright in the Antilles. These included the following four species for 
which original material was found at HAC and for which lectotypification is 
necessary: Gonolobus stephanotrichus Griseb., Orthosia acuminata Griseb., 
Orthosia oblongata Griseb., and Poicilla ovatifolia Griseb. (Table 1). 

Grisebach studied Wright material distributed to him by Asa Gray (GH). 
However, these speci were not necessarily true duplicates of a single gath- 
ering. Indeed, there have been instances of mixed material joined by a single 
number, as Gray appears to have provided the same number to specimens 
Wright may have collected from different locations on different dates (Howard 
1986, 1988). Thus, Howard (1986) stated [brackets ours]: 

“The determination of the type collection, therefore, depends on examin- 
ing the sheet Grisebach saw and named, which is preserved in Géttingen [GOET, 
Universitat Gottingen]. This should match one of the fragments preserved by 
Gray. The GOET specimens however, rarely have field tickets, so the date and 
sored of the type collection can only be determined, if at all, from the GH 
she 


oe (1988) noted that Grisebach likely did not see the Wright collec- 


KRINGS AND FANTZ, 535 


tions sent to the Sauvalle herbarium in Habana (currently deposited in HAC), 
as Asa Gray distributed this material from GH. In contrast to Howard (1988), 
however, it should be noted that the fact that Grisebach may not have seen the 
Wright specimens in the Sauvalle herbarium, has no bearing on their status as 
original material and availability for lectotypification in the future. If judged 
only by this fact, according to Div. 2, Ch.2, Sect. 2, Art. 9 of the International 
Code of Botanical Nomenclature or ICBN (Greuter et al. 2000), they would none- 
theless be considered part of the original material and, as duplicates of one cited 
collection, would be syntypes as articulated by Fantz (1993). 

Gonolobus stephanotrichus.—Two specimens of Wright 2969 reside at 
GOETI. No field tickets accompany the GOET specimens. Both sheets include 
flowers and one also includes a fruit. Fruits are not described in Grisebach’s 
protologue. Duplicates of this number are found at BREM!, G!, GH!, HAC), K!, 
NY!, P!, UC), US!, and reported for MO, although this specimen could not be ob- 
tained on loan. The collection labels of the sheets at GOET are blue and bear 
the dates 1856-7 crossed out. No additional dates are given. The collection la- 
bels for duplicate numbers housed everywhere else, but HAC, are white and 
bear the dates 1860-64 (including the MO specimen, an image of which could 
be seen from the Missouri Botanical Garden website: http://www.mobot.org). 
The collection label for the HAC specimen is white and bears the date 1865. 
This specimen is sterile and quite poor due to insect damage. Field tickets re- 
maining with the sheet at GH seem to indicate that the three mounted frag- 
ments (all in flower) were collected at different times Jan., Mar, Apr.). However, 
it isimpossible to tell which fragment is associated with which field ticket and 
furthermore, none can be correlated with the GOET sheets, as these lack field 
tickets altogether. All material under the number 2969 (incl. the duplicate at 
HAC) does appear to belong to the same species. Thus, in light of the available 
facts, the Wright 2969 sheet at GOET that bears flowers, but lacks fruit, is herein 
designated lectotype for Gonolobus stephanotrichus Griseb. This action agrees 
witha 1984 annotation by R.A. Howard whose lectotypification (1988) appeared 
ina microfiche appendix, not in print, and thus is not effectively published and 
has no standing in nomenclature. Rankin @ Greuter (2000) reported a similar 
case in Antillean Aristolochia. The remaining extant sheets distributed under 
Wright 2969 retain their status as syntypes. An additional fruiting specimen of 
Gonolobus stephanotrichus is housed at S, bearing a Wright label of 1861 and 
the preliminary number 164. Although a determination on the label is provided 
in Grisebach’s hand, we do not consider the specimen original material as fruits 
were not mentioned in the protologue. 

Orthosia.—The only known type material for O. acuminata (Wright 2966) 
is deposited at BM, G, GH, HAC (2 sheets), and K. These specimens bear white 
labels with the dates 1860-64. The mounted field ticket on the GH sheet reads: 


536 BRIT.ORG/SIDA 22(1) 


“Asclepias—Fl. (except the white stigma) green. Farallones San Andre Oct 27.” 
As Wright 2966 (GH) contains fifteen inflorescences and is in very good condi- 
tion; itis here designated as the lectotype for Orthosia acuminata Griseb. Wright 
2966 (G!) contains four inflorescences and is in superior condition to the HAC 
material. Wright 2966 (BMI, G!, HAC!, K!) should be considered isolectotypes. 

Original material of Orthosia oblongata was located at BM!, G!, GH!, HAC, 
K!, and S!, and reported for MO, although the specimen could not be obtained 
on loan. These specimens all bear white labels with the dates 1860-64 (includ- 
ing the MO specimen, an image of which could be seen from the Missouri Bo- 
tanical Garden website). The mounted field ticket of Wright 2967 (GH) reads: 
“Asclepias—Fl. green—a white speck at the tips of the segments. Stigma white. 
Loma de Ranjel June 17.” Field tickets do not accompany the other specimens. 
Wright 2967 (GH) is herein designated lectotype for Orthosia oblongata Griseb., 
considering the duplicates at G, HAC, K, MO (provided that the specimen does 
not constitute another species), and S isolectotypes. Wright 2967 (GH) is in good 
condition, with numerous inflorescences. 

Lectotypification decisions for both Orthosia names were based on the in- 
terpretation that sheets of both Wright 2966 & 2967 are original material, as 
they were respectively cited in the according protologues. There is no evidence 
that they were collected from different localities, although this cannot be ruled 
out altogether, given the notorious problems with Wright collections. However, 
in our opinion, when there is no specific evidence to the contrary, if collection 
number and identification match, the protocol ought to hold that the speci- 
mens be considered duplicates of a single gathering. We recognize that 
“isolectotype” is not an ICBN type designation, but assign it herein should lec- 
totypes become lost or destroyed or additional syntypes are located that we have 
not examined. 

Poicilla ovatifolia.—Syntype material (Wright 2965) was located at BMI, 
BREMI, G!, GH!, GOET!, HAC), K, NY!, P!, S!. and UC! No field tickets accompany 
the GOET specimen or any syntype, except the GH specimen. Accompanying 
field tickets of the GH specimen suggest that the sheet is comprised of at least 
two collections made at different times (Mar, June), although three fragments 
are mounted. The two fragments mounted on the right contain inflorescences; 
the fragment mounted on the left contains infructescences. Both field tickets 
refer to flowers with neither one mentioning fruits. Fruits are not described in 
Grisebach’s protologue and are not present on any other syntype beside the GH 
specimen. Except for the GOET specimen, collection labels of all other known 
specimens are white and bear the dates 1860-64. The GOET specimen bears a 
tan label with a printed date of 1860, although the zero appears to have been 
crossed out. It is heavily written on in Grisebach’s hand and is herein desig- 
nated lectotype for Poicilla ovatifolia Griseb. The studied (and matching) 


duplicately-numbered material in other herbaria remain syntypes. 


KRINGS AND FANTZ, 537 


ACKNOWLEDGMENTS 


We thank the curators and staff of the following herbaria for searching, or pro- 
viding access or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, 
BsG RUE GC, CGE, COLO, CReDUKE 2 ETL PEAS ER PIG .G GHaGOEr1,-H, 
HAC, HAJB, HBG, IA, JJ, ISC, JBSD, JE, K, LD, M, MICH, MIN, MO, MSU, NEU, 
NSW, NY, O, OXE P, PH, RSA, S, U, UC, UPS, US, USF TUR, WILLI, WU, Z. AK 
also thanks Eldis Bécquer and Lutgarda Gonzalez for stimulating conversation 
regarding typitication and extends grateful appreciation for the kind hospital- 
ity afforded by the Jardin Botanico Nacional, Habana, and especially Fabiola 
Areces and Julio Lazcano. Thoughtful reviews of a previous version of the manu- 
script were provided by Carlos Sanchez and Alain Loigier. 


REFERENCES 


Cardenas Atvarez, A.and P. Herrera Ouiver. 1991. Catdlogo de las colecciones histdricas y de 
tipos del Herbario de la Academia de Ciencias de Cuba. Editorial Academia, La Habana. 

FANTZ, PR. 1993. Notes on Clitoria (Leguminosae) in Southeast Asia. Novon 3:352-355. 

GreuTer, W., J. McNelt, FR. Barrie, H.-M. Buroet, V. DEMOULIN, TS. Fitcueiras, DH. Nicotson, P.-C. Sitva, 
J.E. Skoe, P. TREHANE, N.J. TuRLAND, and D.L. Hawkswortu. 2000. International code of botani- 
cal nomenclature (St Louis Code).Regnum Veg. 138. Koeltz Scientific Books, Konigstein 

GriseBach, A.H.R. 1866. Catalogus plantarum cubensium. Wilhelm Engelmann, Leipzig. 

Gutiérrez Amaro, J., |. ARIAS GRANDA, H, Manitz,and E. ALGuiLar CeLpa. 1997.Los tipos del herbario 
“Prof. Dr. Johannes Bisse” del Jardin Botanico Nacional (HAJB): |. Revista Jard. Bot. Nac. 
18:21-50. 

Howard, R.A. 1986. Notes on Quiina (Quiinaceae) and Ilex species (Aquifoliaceae) in Cuba. 
Brittonia 38:13-16. 

Howarp, R.A. 1988. Charles Wright in Cuba, 1856-1867. Chadwick-Healy, Alexandria 

Rankin Rooricuez, R.and W.Greuter. 2000. Notes on Aristolochia linearifolia and A. stenophylla 
(Aristolochiaceae), a vicarious species pair from the Greater Antilles (Cuba and 
Hispaniola). Willdenowia 30:131-139. 


538 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


Joun Dawson and Ros Lucas. 2005. The Nature of Plants: Habitats, Challenges 
& Adaptations. (ISBN 0-88192-675-2, hbk.). Timber Press Inc. 133S.W. Second 
Ave, Suite 450, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, 
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). 
$39.95, 314 pp., 205 color photos, 5 color drawings, glossary, and index, 7.38" 
x 10.38". 

The new book called The Nature of Plants is a plant lover's dream. Have you ever wondered how it’s 

possible for any plants grow in the tundra? Or how underwater plants get their flowers pollinated? 

The authors have provided readers with a wonderful resource to answer these and many of other 


questions about plants surviving in extreme environments. ue textis peppered with rich color pho- 
de 


tographic examples of the plants and | | in the tex 

In each chapter, the reader is introduced to unique environments, conditions, and climates; 
and then told about all the special challenges faced by plants that live there. For example, we're told 
about parasitic plants, desert plants (both hot and cold), fire effects, salt ene plants on windy 


terrain an interactions with animals, fungal and even bacterial interactions between the plants. 
e authors provide a good informational background for the reader regarding the environ- 
mental i climatic situations of the plants. For example, in one chapter titled “Not Enough Water” 


= : 


the authors include an explanation of why deserts even exist and where they're located. This book 
covers a lot more than just North America when they go into detail on the deserts of the world in- 
ee temperate deserts and the special adaptations that plants make for each environment. 
eading about the pee adaptations | to these challenging environments makes this book par- 
aie interesting. After a | lescript { climates or special conditions, the author includes 


ormation about common plant surv ‘val strategies. For example, the chapter about fire shows how 


in 
some plants use fire as a regeneration process; after being damaged, they release seeds or flower. Very 
often, the strategies are given with specific plant examples along with beautiful color photographs 
that help the reader visualize such ceaned fel sicaseas These images are mace wondertul 
because many of the images shov nts i tic locations. In addition, one chapter focuses on the 


various methods of reproduction ne the different vectors for pollen transfer. This sn included 
a wonderlul discussion on the benefits and problems of self{-pollination and out-crossing in plants. 
The book The Nature of Plants: Habitats, Challe nig s & Adaptations is recommended for anyone 


with an interest in plants and their ability t urvi he harshest of habitats. This book does 
not require much botanical background to enjoy, although an understanding of sane family traits 
will be helpful. A glossary is included for the less common terminology. All topics are explained 
with diverse examples and fantastic color photographs. Read The Nature of P “e Habitats, Chal- 
lenges & Adaptations and take a peek into the fascinating survival strategies of plants in extreme 
situations and environmental conditions.—Lee Luckeydoo, Herbarium, Botanical Research Institute 
of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(1): 538, 2006 


COMMENTS ON THE GERBERA-COMPLEX 
(ASTERACEAE: MUTISIEAE) 


Hans V. Hansen 


Botanic Garden 
Ae 2 of Copenhagen 
er Farimagsgade 
Dk . fee oes K, DENMARK 


ABSTRACT 


The recent transfer of Chaptalia hintonii to Gerbera is doubted, as is any transfer of New World 
members to the Old World part of the Gerbera-complex. On a biogeographic basis it seems more 
probable that Gerbera is an Old World clade, while Chaptalia, Trichocline, and T. hieracioides(some- 
times considered to be a Ger per are New Word groups. The geographic distribution of Leibnitzia 

rth America, Asia) and Aml lata (W. Australia) is enigmatic. Additional input of 


ee is needed for a full Spaces of the phylogeny in the Gerbera-complex 


RESUMEN 


f. a Py cal | a baa aad: es Pees ] 


Ur ansfe rencia 
de elementos del BEKO Mundo ala Datte del compl Gerbera del Viejo Mando: En base a la 
] 


biogeografia l Ae: Mune mientras que og eae 
evi 


L t 
Thies ioide  ( | J 


ipachogene 
Mundo. La distribucion geogrfica de Leibnitzia (Norte iva ee. Asia) y Ambbsper ma a 
(Oeste de A tica. Se necesitan datos adicionales para una completa comprension 


b 


de la nee del somiplejs Gerbera. 


In recent papers, Katinas (1998, 2004b) and Nesom (2004a,b) discussed the ge- 
neric position of Chaptalia hintonii Bullock within the Gerbera-complex (the 
scapose complex of Mutisieae subtribe Mutisiinae sensu Cabrera 1977). Their 
discussion continues the long-standing debate regarding OTUs within this com- 
plex. To recapitulate briefly, Jeffrey 967) used LM to point out characters of 
taxonomic value in the group, mainly pertaining to cypselar pubescence and 
pappus-hairs. When [ took up similar studies (e.g., Hansen 1985, 1990) SEM had 
been introduced, but my conclusions deviated little from those of Jeffrey. 

Nesom (1983) contributed with his first study of the Gerbera-complex 
(American Leibnitzia) and next focused on Chaptalia Vent. (Nesom 1984a,b, 
1995, 2004a,b; Cabrera & Nesom 2003), while other workers have also published 
studies dealing with the scapose group (Katinas 1998, 2004a,b; Sancho & 
Katinas 2002; Moraes 1998; Hind 1999, 2001; Roque 2005). All these studies, with 
their general access to living material, have amplified our knowledge about the 
Gerbera-complex. 

The complex includes about 100 species, and evidently it takes subtle char- 


SIDA 22(1): 539 — 543. 2006 


540 BRIT.ORG/SIDA 22(1) 


acters to delimit monophyletic groups within it. This is noteworthy in view of 
its wide distribution: Trichocline Cass. Lulia Zardini, and Chaptalia Vent. in the 
New World, Leibnitzia Cass. in North America and Asia, Gerbera L. in Africa, 
Madagascar, and Asia (i.e., Gerbera sensu Hansen 1990), Perdicium L. in W. Cape, 
Uechtritzia Freyn in Asia,and Amblysperma spathulata (A.Cunn.ex DC.) DJ.N. 
Hind in W. Australia. It has been disputed whether Amblysperma falls within 
the limits of Trichocline (Hind 2001, and implicitly Hansen 1990, contra Zardini 
1975 and Katinas 2004a), and whether the transfer of Trichocline hieracioides 
(Kunth) Ferreyra (Ecuador, Peru) to Gerbera by Zardini (1974) is correct. The 
most significant problem, however, is with Chaptalia, for which no complete 
revisionary treatment exists; Burkart (1944) only considered part of the genus 
in detail. 

Nesom (2004a) and Katinas (2004b) remarked that while Jeffrey would 
split the complex into smaller genera, | suggested the recognition of one large 
genus (hence Gerbera). This calls for an explanation. The relatively ancestral 
position within Asteraceae of Mutisieae (however circumscribed, but at least 
excluding subfamily Barnadesioideae (Benth. G Hook.f.) K. Bremer & R.K. 
Jansen) is now supported by morphologic and molecular evidence. Of relevance 
here is that part of the classic Mutisiinae in all studies is indicated to be mono- 
phyletic, with the Gerbera-complex placed in a relatively advanced position. 
Actually, the Brazilian monotypic genus Lulia (Zardini 1980) (ie., Trichocline 
nervosa Less.), by its monocephalous, albeit non-scapose habit, stands between 
the Gerbera-complex and its sister (a group including Mutisia Lf, Chaetanthera 
Ruiz & Pav, Duidaea S.F Blake, and Pachylaena D. Don in Kim et al. 2002; Lulia 
was not considered). 

In Hansen (1991), | anticipated these aspects and then reasoned that if 
Mutisia and Chaetanthera are not split into minor taxonomic units, it would 
not be logical to split the Gerbera-complex either, hence all species should fall 
within Gerbera. This explains my way of reasoning which was not versus Jef- 
[rey (1967), but simply an alternative provided by cladistic reasoning. 

Katinas (2004b:938-939) presented a key to the scapose group with seven 
genera (including Lulia, but with Amblysperma sunk into Trichocline and 
awaiting a complete revision of Chaptalia). In this key there were 
autapomorphies for each genus (if we accept that Trichocline hasa special type 
of cypsela hairs). However, there is a complication in Katinas’s key couplet 6, 
since Gerbera p.p. (sects. Gerbera, Parva, and Isanthus) do not have the 
apomorphic state ‘trimorphic rays. This is precisely where the problems with 
Gerbera (in its current sense) arise. 

Nesom (2004a, cl. Nesom 1995; Cabrera & Nesom 2003) has maintained 
that Chaptalia hintonii belongs to Chaptalia sect. Chaptalia. The traits which 
led Katinas to move it to Gerbera (all florets bilabiate, outer ray florets long, 
ray florets with staminodes) can be found in various Chaptalia, and the cypsela 


HANSEN, COMMENTS ON THE GERBERA-COMPLEX 541 


hairs in C. hintonii do not deviate in morphology from those in Chaptalia sect. 
Chaptalia p.p. (the C. lyratifolia group). 

I believe that Nesom has the strongest point, namely that the ‘deviating 
traits in C. hintonii do not disrupt its alliance with Chaptalia sect. Chaptalia. 
Rather, C. hintonii possesses states which are parallelisms/reversals to those in 
Gerbera. As Nesom indirectly argues, it seems more probable on a biogeographic 
basis that Gerbera is an Old World clade, while Chaptalia, Trichocline, and the 
enigmatic T. hieracioides are New World groups (not overlooking problems with 
Amblysperma in W. Australia and Leibnitzia, which is divided between Asia 
and North America). 

Evidently, various authors during their close survey of taxa detect excep- 
tions to the currently used taxonomy and thus find reason to allocate this or 
that species. But the question is whether we push the case too far. We know 
more today than we did in Hansen (1990) and hardly disagree on the polariza- 
tion of states. Hence, a matrix with all relevant placeholders in the Gerbera- 
complex treated as OTUs ultimately should be prepared, because only then it 
will be revealed, if the addition of new characters can disrupt the unity of (some 
of) the currently recognized taxonomic groups. Presumably, however, the con- 
sensus tree will still be burdened with polytomies, since the number of known 
synapomorphies is so small. Additional input of data is therefore much wel- 
come. However, along with the detection of amplified variation in ray floret 
morphology, cypselar vestiture, etc. the selection of relevant characters may 
cause more and more debate. A good example is the presence of trichomes on 
corollas in 13 species of Trichocline, three of Onoseris, and one investigated spe- 
cies of Uechtritzia (Sancho & Katinas 2002). Is this information of phyloge- 
netic relevance or not? 

Nesom (2004a:932) has now suggested that Chaptalia sects. Lieberkuhna 
(Cass.) Burkart and Loxodon (Cass.) Burkart possibly may be separated at ge- 
neric rank, among other things due to their distinctive cypselar vestiture. 
Katinas (2004b) likewise predicted taxonomic alterations at generic rank. The 
discussion between the two authors reveals considerable disagreement, both 
with respect to observation of characters (e.g., whether ray flower staminodes 
are present or not) and how to interpret their phylogenetic relevance. | must 
stay neutral, since I never studied Chaptalia in detail, but clearly the main is- 
sue is to decide whether Chaptalia is monophyletic (even if split into two gen- 
era), with evolution of character states parallel to those in Gerbera, or whether 
some species of Chaptalia really should be moved to Gerbera or even some- 
thing else. 


? 


ACKNOWLEDGMENTS 


I thank Guy Nesom and an anonymous reviewer for their most insightful com- 
ments which improved much on this paper. 


542 BRIT.ORG/SIDA 22(1) 


REFERENCES 


Burkart, A. 1944. Estudio del género de Compuestas Chaptalia con especial referencia a 
las especies argentinas. Darwiniana 6:505-594, 

Caprera, A.L.1977.Mutisieae-systematic review.|n: Heywood,V.H.,J. Harborne, and B.L.Turner, 
eds. The biology and chemistry of the Compositae I]. Academic Press, London. Pp. 
1039-1066. 

Caprera, L.and G.L. Nesom. 2003. A new species of Chaptalia (Asteraceae: Mutisieae) from 
Mexico and rediscovery of Chaptalia mexicana. Sida 20:1363-1369. 

Hansen, H.V. 1985. A taxonomic revision of the genus Gerbera (Compositae-Mutisieae) 
sections Gerbera, Parva, Piloselloides (in Africa), and Lasiopus. Opera Bot. 78:1-36 

Hansen, H.V. 1990. Phylogenetic studies in the Gerbera-complex (Compositae, tribe 
Mutisieae, subtribe Mutisiinae). Nordic J. Bot. 9:469-485. 

Hansen, H.V. 1991. Phylogenetic studies in Compositae tribe Mutisieae. Opera Bot. 109: 


Hino, DJ.N. 1999. Notes on Chaptalia (Compositae: Mutisieae) in Brazil. Kew Bull. 54: 
933-939, 

Hino, D.J.N.2001.A new combination in Amblysperma (Compositae: Mutisieae). Kew Bull. 
56:711-713. 

JerFrey, C. 1967. Notes on Compositae: II. The Mutisieae in east tropical Africa. Kew Bull. 
21:177-233. 

Karinas, L. 1998. The Mexican Chaptalia hintonii is a Gerbera (Asteraceae, Mutisieae). Novon 
8:380-385. 

Karinas, L.2004a.Amblysperma should be retained under Trichocline (Asteraceae, Mutisieae) 
Taxon 53:108-112. 

Katinas, L. 2004b. The Gerbera complex (Asteraceae: Mutisieae): to split or not to split. Sida 
21:935-940. 

Kim, H.G., D.J. LoockerMAN, and R.K. Jansen. 2002. Systematic implications of ndhF sequence 
variation in the Mutisieae (Asteraceae). Syst.Bot. 27:598-609. 

Moraes, M.D. be. 1998. Chaptalia hermogenis (Asteraceae: Mutisieae), a new species from 
the Brazilian Atlantic Rain Forest. Novon 8:173-175. 

Nesom, G.L. 1983. Biology and taxonomy of American Leibnitzia (Asteraceae, Mutisieae). 
Brittonia 35:126-139. 

Nesom, G.L. 1984a. A new, widespread species of Chaptalia (Asteraceae: Mutisieae) from 
Mexico. Rhodora 86:127-130, 

Nesom, G.L. 1984b. Taxonomy and distribution of Chaptalia dentata and C. albicans 
(Asteraceae: Mutisieae). Brittonia 36:396-401. 

Nesom, G.L. 1995. Revision of Chaptalia (Asteraceae: Mutisieae) from North America and 
continental Central America. Phytologia 78:153-188. 

Nesom, G.L. 2004a. Generic placement of Chaptalia hintonii (Asteraceae: Mutisieae). Sida 
21:929-933. 

Nesom, G.L. 2004b. Response to “The Gerbera complex (Asteraceae: Mutisieae): to split or 
not to split” by Liliana Katinas. Sida 21:941-942. 


— 


HANSEN, COMMENTS ON THE GERBERA-COMPLEX 543 


Roque, N. 2005. A new species of Chaptalia (Compositae, Mutisieae) from Minas Gerais, 
Brazil. Kew Bull.60:133-135. 

SancHo, G. and L. Katinas, 2002. Are the trichomes in corollas of Mutisieae (Asteraceae) 
really twin hairs? Bot. J. Linn. Soc. 140:427-433. 

ZaARDINI, EM. 1974. Sobre la presencia del género Gerbera en America. Bol.Soc. Argent. Bot. 
16:103-108. 

ZARDINI, E.M. 1975. Revisi6n del género Trichocline (Compositae). Darwiniana 19:618-733. 

ZARDINI, E.M. 1980. Lulia - un nuevo genero de Compuestas. Bol. Soc. Argent. Bot. 19: 
255-258. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


He_en Heipert. 2000. Papermaking with Garden Plants and Common Weeds. 
(Paperback edition). (SBN 1-58017-622-4, pbk.). Storey Publishing, 210 MASS 
MoCA Way, North Adams, MA, 01247, U.S.A. (Orders: # 67622, 1-800-441- 
5700, www.storey.com ). $14.95, 112 pp., color photos, drawings, appendix, 
glossary, resource guide, index, 91/4" x 10" 

Papermaking With Garden Plants And Common Weeds isa good resource for crafters and profession- 

als interested in making their own paper from natural materials. Author Helen Heibert has put to- 

gether a wonderful reference with ee instructions on three methods of papermaking with 

a variety of natural materials. These instructions include practical advice on keeping records of re- 

sults with different materials, detailed fiber processing instructions, and sidebars of personal expe- 


riences and successes of the author. 

he book begins with preliminary information on responsible and safe plant fiber and mate- 
rial collecting. This eee information is followed by descriptions of the types of possible 
fibers: bast, leaves, grass and ees materials oe uae ad and recycle fibers. Next she pre- 


f king four mould iia cee a 
tep-by- 


sents the 
with the different ee techniques. a the different alkali recipes, she presents 
ssing instructions covering cooking, beating, retting, and safety information. In a 
S Wit 


step proce 
to that, she includes information on the additives for protecting and coloring fiber materia 


> 
=) 


artificial and natural dyes. 
Along with the Co instructions, the author has ae included great color images that 


highlight the step-by | the three Western style with a mould 


and deckle, Japanese paper eels with a sugeta and formation aid (recipes included), and t 
There is a helpful troubleshooting section for potential sian 


os 


e Ne- 


palese method using a deckle bo 
that may be encountered Pee the paper making process. Multiple pressing techniques are also 
presented with color images and include instructions on how to make your own paper press as well 
as wet paper handling suggestions and drying technique 

e author has included prone with instructions ol ten different artists who used ten differ- 
.A small personal story box is included with 


ent plant materials and pa 
each artist's project agattionahy, seven step- ry oe projects are presented including: flower petal 
envelope, lampshade, and accordion book 


within paper, | 
ealowing the Section on projects is important iaigr mation including lists of papermaking plants, 


with common and Latin names, glossary, reading list, and a resource guide 


Author Helen Heibert has provided adventurous crafters with well-guided instructions on 
paper- making techniques and projects. The et are recommended for adults and, due to the 
hemical king involved, children with dervision. The paper types and projects sug- 


gested and included in the book Papermaking with Garden Plants and Common Weeds are promising 

for scra BOSanE a making, craft groups, interior design, education, youth projects and man 
8 8 y 

_uckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pe- 


othe 
can cee Fort Worth, TX 161 “ 4060, U.S.A. 


SIDA 22(1): 544, 2006 


VALIDATION OF THE NAME DAHLIA SUBLIGNOSA 


(ASTERACEAE) 
Dayle E. Saar Paul D.S@rensen 
Department of Biological Sciences Department of Biological Sciences 
Murray State University Northern Illinois University 
Aurray, Kentucky 42071, U.S.A Dekalb, Illinois 60115, U.S.A. 


Article 33.3 of the International Code of Botanical Nomenclature (ICBN) (St. 
Louis Code 2000) states that a new combination based ona previously and val- 
idly published name is not itself validly published unless its basionym is clearly 
indicated and fully referenced. While the original work was cited regarding the 
new combination of Dahlia dissecta S. Watson var. sublignosa P.D. Sorensen to 
Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, the basionym was 
not “clearly identified” (Saar & Sorensen 2005). Therefore, the combination is 
validated here in accordance with the ICBN. 


Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, comb. & stat. nov. 
BASIONYM: Dahlia dissecta var. sublignosa PD. Sorensen, Rhodora 71:35]. 1969. TyPE: MEXICO. 
Tamaulipas: 4.8 km N of Miquihuana in forest dominated by Pinus (99°47' N Lat; 23°36' W 
Long.), elev. ca. 2100 m, 14 Jul 1949, Stanford, Taylor, & Lauber 2436 (HOLOTYPE: NY; ISOTYPES: 
GH, MICH, TEX, UC, US-2, WTU). 


Asan explanatory note, the holotype specimen collected by Stanford, Taylor, & 
Lauber 2436 cited in our paper is a correction of the original basionym, which 
was incorrectly stated as collected by Stanford, Rutherford, and Northcroft 2436. 
ACKNOWLEDGMENTS 
We thank Teri Bilsborrow for bringing this oversight to out attention and Kanchi 
Gandhi for helpful comments regarding this validation notice. 
REFERENCES 


Saar, D.E.and PD. Sorensen. 2005. Dahlia subli pecies in its own right. Sida 21:2161- 
2167. 


SIDA 22(1): 545. 2006 


546 BRIT.ORG/SIDA 22 


— 


1) 


Book REVIEW 


FREDERICK R. TROFH and Louis M. THompson. 2005. Soils and Soil Fertility. 6th edition. 
(ISBN 0-8138-0995-X, hbk.). Blackwell Publishing. 2121 State Ave., Ames, 

TA 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. 
one. 515-292-0140, 515-292-3348 fax 1-800-862-6657, www.blackwell 

» Vix | (7) LI] ] lly 1 ae lee Pee g.com). $89.95, 498 pp., 


231 ‘iiietaciona. 7 210" 


understood lan- 


Soil and Soil Fertility is a well-illustrated book that presents soil t 
guage. It would be suitable as a textbook for classes on general soils or soil fertility. Additionally, this 


title could also be a beneficial reference text for classes in agronomy, hydrology and agricultural 


engineering. 
Within the preface, the authors mention that this book has been “designed for use as a textbook 
for the introductory course in soils for students in agriculture and related sciences.” Chapter topics 
include 1) Soil; 2) Soil Formation; 3) Physical Properties of Soils; 4) Soil Mechanics; 5) Soil Water; 6) 
Soil Organic Matter; 7) Soil Mineralogy; 8) Soil Chemistry; 9) Amending the Soil; 10) Fertilizers; 11) 
Nitrogen; 12) Phosphorus; 13) Potassium; 14) Calcium, Magnesium, and Sulfur; 15) The Micronutri- 
ents, 16) Variations in Plant Composition; 17) Soil Classification; 18) Land Use and Soil Manage- 
ment; 19) Water Management, 20) Soil Erosion and Its Control; and 21) Soil Pollution. The order of 
chapters How well into one ae pune on previous chapters’ materials, starting with soils 


basics and mineralogy, d g and fertility and ending with environmental topics 


and land stewandehip 
The text is very readable and is accompanied by many supportive illustrations, cule and 
charts. These diagrams and een on Lae ize ee ene white wee mans ne other e easily dis- 
tinguishable designs. TI I lly offer an 
example problem with éeplonation: Basic soil tests/analyses are iced hg the text in- 


— 


cluding items such as particle size analysis using Stoke’s law, a hydrometer, and a RUSLE soil loss 

equation example. At the end of each chapter is an im et notes section, and a ” of questions 
that if answered, would demonstrate an understanding of the text 

In addition to all of the discussion of general soils information, this book 

other pn tant souls Subiccts:> soil cea and soil classification. The section on soil rie isa 

ehel il fertility fllowee Hace daunaaors nou 

s in plants, the 


] Jp] einr 


ee micronutrients. These chapters include chemical use and deficiency symp 
nutrient cycle and availability, fertilizer types for that nutrient, application info “e more. The chap- 
ter on soil classification includes a number of different systems with background and explanation 
on each system, additionally covering land use and land practices to reduce soil loss. In the end, 


there isa measurement and con Nees summary as well as a glossary. 
Soil and Soil Fertility by Troeh and Thompson would be an excellent textbook for soils and 
related classes. oS isa —— intelligent book os l of s sup por tive black and w pied illustrations within 


xt. Althou Sa 
the : gh g textbook g ng this text for 


=e mation and a better understanding of soil properties and soil nese ee oa: keydoo, Her- 


SIDA 22(1): 546. 2006 


A NEW COMBINATION IN POLYTRICHASTRUM 
(BRYOPHYTA: POLYTRICHACEAE) 


G.L.Smith Merrill 
Department of Botany 
The Field Museum 
1400 South Lake Shore Drive 
res ey linois 60655-2496, U.S.A. 


ABSTRACT 


(NA : \ 


The new combination Polytrichastrum alpinum var. sylvaticum L. Merrill is provided. 


RESUMEN 


L 


Se aporta la nueva combinacion Polytrichastrum alpinum var. sylvaticum (Menzies) GL. Merrill. 


The following transfer is proposed for use in a treatment of the moss family 
Polytrichaceae in the Flora of North America North of Mexico, vol. 27, the first 
of the projected three bryophyte volumes. Additional discussion will be found 
in the flora treatment. 


= 


Polytrichastrum alpinum var. sylvaticum (Menzies) G.L. Merrill, comb. nov. 
BASIONYM: Polytrichum sylvaticum Menzies, Bot. Zeit. (Regensburg) 1:74. 1802. TyPE: U.S.A: 
Northwest Coast of North America, Menzies (HOLOTYPE: E!). 

Pogonatum alpinum var. aye Scien Lindberg, Bot. Not. 1867: 130. 1867. Polytrichum 
alpinum var. sylvaticun ndberg, Acta Soc. Sci. Fenn. 10:6. 1872, “silvaticum.” 
Polytrichum macounti iment in ene Bull. Torrey Bot. a 16:96. 1889; Pogonatum 

macounii (Kindberg) Kindberg & Macoun, Cat. Canad. Pl. 6:152. 1892. Pogonatum alpinum 
var. macounii (Kindberg) Cardot & Thériot, Proc. Wash. Aon Sci. 4:328. 1902. Polytrichum 
alpinum var. macounii (Kindberg) Saito, J. Jap. Bot. 31:75. 1956. Pogonatum alpinum subsp. 
macounii (Kindberg) Persson in Persson & Viereck, Lindbergia 9:16. 1983 


Habitat and distri onen in North America.—On soil in moist woods, common in the 
lowlands, iddl British Columbia, southern Alaska, Oregon, Washington. 

Polytrichastrum alpinum i is a widespread, polymorphic species, common 
at middle to upper elevations throughout the North Temperate zone, in south 
temperate South America, the Antarctic, and Australasia. The var. sylvaticum 
isa giant form of the Pacific Northwest with stems as muchas 14 cm high, simple 
to laxly branched, sharply serrate leaves to 19 mm long, and large, plump, cy- 
lindric capsules, often somewhat zygomorphic and widest below the middle. 


REFERENCES 
Frye, T.C.1937. Polytrichaceae. In: Grout, AJ. Moss flora of North America north of Mexico. 


Pp. 99-128 
Lawton, E.1971.Moss flora of the Pacific Northwest. Hattori Botanical Laboratory, Nichinan,Japan. 


SIDA 22(1): 547, 2006 


548 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


STEVEN CARROLL and STEVEN SALT. 2004. Ecology for Gardeners. (ISBN: 0-88192- 
611-6, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 
Oi 204-5021. US.A.(Orders: www.tim berpress.com, il@timberpress.com, 
503-227-2878, 1-800-327-5680, 503-227-3070 fax). $29.95, 420 pp., ll baw 
figures, 170 color photos, glossary, and common, scientific and subject 
indices, 6" x 9" 


=) 


With Ecology For Gardeners, authors Steven Carroll and Steven Salt have written a book that is es- 


= 


sential for any gardener’s shelf. Their book keeps ecological principles in mind while dealing with 


the subject of gardening. 


Ecology for Gansenirs provide x ii many? ints to pau 1 think about. It contains 


many cause & effect situations anc between abiotic and biotic factors in 


rs 
your garden. The text includes discussion of seemingly Go topics that one may not instinc- 


tively link together at first; such as how soil temperature relates to nutrient availability for plants, 
intraspecific competition and plant spacing, pruning as herbivory, energy roles in a garden, the abil- 


ity of the wind to alter the shape a garden, the importance of microbes to the nutrient cycling & 
avai a ity in oe puch etc. neler topics include plant biology, non-plant aan ai 
heir environment ts,an 


oo 


gi — as applied a (stewardship and gar i management) 
Each chapter begins with a basic and easy to understand intro dueion about the topic. After 
the introduction ios each ¢ Baptett ane fopies is div ute into were organized es For example, 


the chapter i etition, herbivory, pre- 


oO 
pa 
Se) 


dation, parasitism and mautualism, Another example is the garden environment chapter that is di- 


Vv ee into sun, air, water sell energy, food isha and nutrient Se within the alee: - stem. The 


tyle is 


the chapter on appliec eee in etn ane isa subtopic of ateee management. I con- 


- 


tail cussion on mulching varieties, benef 


each mulch type. The chapter 


on applied ecology in the garden really helps th the reader bring together all the previous information 


of other chapters and may lead to many “Eureka!” moments. Clear color ee phs exemplify many 
ach 


of the various plants, items, and occurrences referenced within e 


Ap 


The book ee logy, for Garde ners is full many a provoking topics; especially for com- 
i nd E 


monly used gical ba hind them. The authors have done a fan- 
tastic job of ever ts eedlonics! part of gardening very accessible to gardeners. Readers will defi- 
nitely understand more of the ecology that occurs in, uae and pada the plants in their garden. 


More importantly, gardeners will be able to make more informed garden management decisions af- 
ter reading Carroll and Salt’s Ecology for Gardeners —Lee Lucke ydoo, Herbarium, Botanical Research 
Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(1):, ): 548. 2006 


A NEW COMBINATION IN TETRAPHIS 
(BRYOPHYTA: TETRAPHIDACEAE) 


Judith A. Harpel 


Herbarium, Burke Museum 
Box 35325, University of Washington 
Seattle Washington 98195-5325, U.S.A. 
wjharpel@earthlink.net 


ABSTRACT 


n Tetraphis pellucida Hedw. var trachypoda 1 | larify the taxonomic 


The mbinat : 


ane - ae moss Reais trachypoda and to recognize the morphological difference of having a 
straight, slightly papillose seta. A revised key to the genus, a diagnosis of var. trachypoda, and speci- 


mens examined are provided. 


RESUMEN 
Se propone la nueva combinacion Tetraphis pellucida Hedw. var. trachypoda, para aclarar el rango 
taxonomico del musgo Tetraphis ee y para reconocer las diferencias morfologicas de tener 
una seda derecha ligeramente papilosa. Se ofrece una clave revisada del género, una ee de la 
var. trachypoda, y los especimenes os 


Kindberg (1893) described Georgia trachypoda from specimens collected by 
John Macoun in British Columbia as follows: “leaves subovate or ovate-oblong, 
short-acuminate and acute. Perichaetial leaves with long, sublinear, acute acu- 
men; costa not excurrent. Capsule straight; pedicel straight, rough in the upper 
part.” In 1900 Paris accepted Hedwig’s genus Tetraphis and Georgia trachypoda 
Kindb. became T. trachypoda (Kindb.) Par. Currently Tetraphis consists of two 
recognized species, neemicwar Cue ex Milde and T. pellucida Hedw. Because 
the gametophytes are undisti ble, these two species are separated by se- 
tae characteristics. Since its description, Tetraphis trachypoda has been recog- 
nized at various taxonomic ranks. 

In 1936 Grout combined G. trachypoda under T. geniculata noting the seta 
roughness and dismissing the straight seta. He comments that “I: geniculata is 
not materially different in macroscopic appearance from T. pellucida, as the 
seta is not always noticeably bent at first glance and the bend is sometimes 
absent as in Kindberg’s Georgia trachypoda (the only difference of importance 
that he notes in his description). Specimens in the herbarium of the National 
Museum of Canada, labeled Georgia trachypoda and collected at same date and 
place as the type and probably a part of it, have the seta only slightly rough- 
ened and not geniculate, otherwise normal.” 

Van Der Wijk et al (1969) and Savicz-Ljubitzkaja and Smirnova (1970) fol- 
lowed Grout’s lead and also placed T. trachypoda under T.geniculata. In Russia, 


SIDA 22(1): 549 — 554. 2006 


550 BRIT.ORG/SIDA 22(1) 


Abramova et al. (1954) regarded T. trachypoda as having only a weakly 
mamillose stalk, and therefore combined it under T. pellucida. They did not 
comment on its distribution in Russia. Crum and Anderson (1981) in their treat- 
ment of Tetraphis, comment, “Rare specimens with some setae more or less 
roughened above but not at all geniculate qualify as Georgia trachypoda Kindb., 
which Grout placed in synonymy of Tetraphis geniculata. The sporadic occur- 
rence of such plants (in Ontario, Minnesota, Montana and British Columbia at 
least) in close association with plants with smooth setae seems to indicate that 
they are anomalous forms of T. pellucida.” Ignatov and Afonina (1992) in their 
check-list of mosses of the former USSR also treated T. trachypoda asa synomen 
of T. pellucida. 

Forman (1962) in his discussion of the family Tetraphidaceae makes no 
mention of T. trachypoda but does comment that “separating the two species of 
Tetraphis is easy when mature sporophytes are present because very few colo- 
nies show intermediate sporphytic characters. In such intermediate cases the 
tuberculate character of the seta is more constant than the geniculate one.” Be- 
tween 1936 and 1992 T. trachypoda has been synonymized under both T. 
geniculata and T. pellucida. 

While examining specimens of Tetraphis from North America for the Bryo- 
phyte Flora of North America project, it became apparent that an additional 
morphological feature of the seta could be used to help clarify the confusion 
surrounding [. trachypoda. Although Weber and Simone (1977) determined that 
the papillose or tuberculate surface of the seta in T. geniculata was the result of 
protruding end-walls of the cells, they did not mention the spiral vs. straight 
arrangement of the cells. Their paper focuses on the nature of the papillae and 
does not comment on the fact that the papillae are also the result of the seta 
cells becoming straight rather than spirally twisted. 

Tetraphis pellucida hasa straight smooth seta, with all of the cells strongly 
spirally twisted throughout its length and it is never papillose. The cells of the 
seta in Tetraphis geniculata are spirally twisted up to the point where the bend 
occurs. Above the bend the cells are straight and conspicuously papillose by 
projecting cell-end walls. Tetraphis trachypoda on the other hand hasa straight 
seta, in which the cells are spirally arranged and interspersed with short sec- 
tions of cells (or sometimes only a few cells) that are straight, which results ina 
few papillae on a normally smooth seta. When the type specimen of T. 
trachypoda was examined, this new characteristic was verified. 

The change of cell arrangement correlated with papillae and a bend in the 
seta of T’ geniculata, and straight, consistently spirally arranged, smooth seta 
cells in T. pellucida appear to be characteristics sufficient to maintain both of 
these taxa as individual species. Tetraphis trachypoda on the other hand ap- 
pears to be intermediate between T. geniculata and T. pellucida, but its charac- 


HARPEL, A NEW COMBINATION IN TETRAPHIS 551 


teristics are not sufficient to maintain it as a separate species. It is suggested, 
therefore, that T. trachypoda be treated as a variety of T. pellucida based on the 
straight, slightly papillose setae, the outermost cells of which have alternating 
zones of cells with either spiraled or straight side walls. 


Tetraphis pellucida Hedw. var. trachypoda (Kindb. ex Paris) Harpel, comb. nov. 
Figs. 1, 2). Georgia trachypoda Kindb., Rev. Bryol. 20:93. 1893. TyPE: CANADA. BRITISH 
COLUMBIA: Sicamous, 7 Apr 1889, J. Macoun s.n. (HOLOTYPE: S!). 


Plants in dense turfs or scattered patches, green above, reddish-brown below. 
Stems 8-15 mm uptolcm tall. Leaves erect-spreading, ovate to ovate-lanceolate, 
1-2 mm long, keeled, costa subpercurrent, margins entire, plane to broadly re- 
curved, upper medial cells smooth, irregularly rounded-hexagonal, lower basal 
cells oblong-linear. Stalked, discoid gemmae present in a rosette of rounded 
bracts (occasionally weakly developed) formed on top of an attenuated stem. 
Seta 6-14 mm long, erect + flexuose, cells lightly papillose usually near the base, 
cells spirally twisted but interspersed with sections of straight cells, twisted 
when dry. Capsule narrowly cylindrical, 2-3 mm long, peristome teeth 4. Spores 
10-17 um, green to yellowish-green, papillose. Capsules mature in spring to early 
summer, rare on well rotted wood, stumps and logs. 

The distribution of T: pellucida var. trachypoda is interesting because it 
occurs in North America and Russia, although at the present time, the geo- 
graphic range in Russia is unknown. Based on the North American material 
examined, it appears to only be found in areas where both T. pellucida and T. 
geniculata occur, which suggests that it would occur in the Russian Far East 
where both of these species also overlap. Examination of material frorn this 
area and additional fieldwork would =e to validate this hypothesis. 


ANADA. British Columbi 7 Apr 1889, Macoun s.n.(S). LABRADOR: 
eras. Bay, 10 Jun 1949, Schofield 32 (NY, UBC). NEW BRUNSWICK. Restigouche Co.: Mount Carleton 
Provincial Park, 12 Aug 1988, Bristow s.n. (MICH). Northwest Territories. Nahanni National Park, 7 Jul 
1974, Scotter 22554: 15 Jul 1976, Steere 76-786, 76-822: 25 Jul 1974, Marsh 4550 (NY). UNITED STATES. 
COLORADO. Gunnison Co.: 3 Aug 1969, Weber B-34769 (COLO, NY). MONTANA. Lake Co.: Flathead 
Lake, 26 Jun 1960, Schofield 11632 (NY, UBC). WYOMING. Yellowstone National Park, 17 Aug 1953, 
Welch 16435 (NY). Teton Co.: 29 Aug 1973, Hermann 25571 (NY) 


KEY TO SPECIES OF TETRAPHIS 


. Seta geniculate, cells smooth, and spirally twisted below the bend, cells straight 
and papillose by projecting cell walls above the bend 1.T. geniculata 
. Seta not geniculate, outer cells smooth, and spirally twisted throughout the entire 
length of the seta, not papillose, or sometimes lightly papillose near the 
2. Seta smooth, superficial cells spirally twisted throughout the entire ae gth 
2a.T. salted: var. pellucida 
2. Seta usually lightly papillose near the base, superficial cells spill twisted but 
interspersed with sections of straight cells 2b.T. pellucida var. trachypoda 


— 


— 


Vow @ene 


Piura sy 
_ at Ff i Trak spats 
ABE 


areas es 
Ob rvs, Cy, Lthiroker 
v9 F9 


eA 


ff 


SRK CK dope 


{cr 


] 


gy 
7 - PP « 


_J.Macoun s.n. (S). Size marker in lower right corner equals 2.5 cm. 


Re oat 
Oh. mr "3 ye, A 
a 
Fb Lite gewigh 


ma 
ed 


L wcoitth 


: : ft sarha 


, de - 


BRIT.ORG/SIDA 22(1) 


HARPEL, A NEW COMBINATION IN TETRAPHIS 553 


AMOS 
var.trachypoda 
\ 


AS) 5 
NP 


© 


CE 
e) 
CO 

STR 


ZO 
JO 


" 4 
ea 


we 
C 
PD 
© 
CY 
<A 


aS ct 
OS: 
© 


Cw 
e) 
zo! 


Se 
Cn 


SF 
OPS 
OAL 


Fic. 2.A. habit 7. pellucid llucid d 7. pellucd h | B. capsules. C. 7. Hucid 


b I YP b 
lls.D gemmae cup E. qemma.F. leaves. G. basal cells. H. upper cells. | Tp lucid 
p.E.g PI 
aera 1 F 


J. habit 7. geniculata K.T.g I 
iation, FNA Vol. 27, artist PM. Eckel 


ACKNOWLEDGMENTS 
I would like to thank the curators and staff from the following herbaria for the 
loan of material used in this treatment, University of Michigan (MICH), New 
York Botanic Garden (NY), Oregon State University (OSU), and Swedish Mu- 
seum of Natural History (S), University of British Columbia (UBC), University 


554 BRIT.ORG/SIDA 22(1) 


of Washington (WTU), and Yellowstone National Park (YELLO). Appreciation 
is extended to Wilf Schofield for his support during the preparation of this treat- 
ment and to Patricia Eckel for her excellent drawing. Barney Lipscomb, Norton 
Miller and William Weber are thanked for providing helpful suggestions in their 
reviews of the manuscript. 


REFERENCES 


ABRAMOVA, A., K. LADYZHENSKAJA, aNd L. Savicz-LuusitzkaJa. 1954. Flora plantarum cryptogamarum 
URSS Vol. Ill Musci Frondosi (2). Typis Et Impensis Academiae Scientiarum URSS Mos- 
cow (in Russian). 

Crum, H.and L. ANDERSON. 1981. Mosses of eastern North America. Vol. 2. Columbia Univer- 
sity Press, NY. 

Forman, R.1962.The family Tetraphidaceae in North America:conti | distribution and 
ecology. Bryologist 65:280-285. 

Grout, A. 1936. Moss flora of North America north of Mexico, Vol. 1. Newfane, VT. 

Icnatov, M.and O. Aronina. 1992. Check-list of mosses of the former USSR. Arctoa 1:1-85, 

KinDBerG, N. 1893. Georgia pellucida et les especes alliees. Rev. Bryol. 20:93. 

Paris, E.G. 1900. Index bryologicus sive enumeratio muscorum huscusque cognitorum 
adjunctis synonymia distributioneque geographica locupletissimis : supplementum 
primum. P.318. 

Savicz-LyusitzkAa, L.and Z. Smirnova. 1970. The handbook of the mosses of the U.S.S.R. The 
mosses Acrocarpous. The Academy of Sciences of the U.S.S.R. The Komarov Botanical 
Institute (in Russian). 

VAN Der Wut, R., W. Marcadant, and P. FlorscHutz. 1969. Index muscorum.Vol.5 (T-Z, Appen- 
dix). Kemink en Zoon N.V. Utrecht. 

Weeer, W.A. and L.D. Simone. 1977. Tetraphis pellucida and T. genciculata: Scindulae as diag- 
nostic features in bryophytes. Bryologist 80:164—-167, 


A NEW COMBINATION IN THE BROMUS CATHARTICUS 
COMPLEX (POACEAE: BROMEAE SECT. CERATOCHLOA) 


Ana Maria Planchuelo 


Herbario ACOR 
Facultad de Ciencias Agropecuarias 
Universidad Naci enel Cordoba 
CONICET, Casilla de Correo 509 
5000 Cérdoba, ARGENTINA 


ABSTRACT 


=) 


A new combination, Bromus catharticus var. elata (E. Desv.) Planchuelo is proposed. This taxon has 
hitherto been treated either as B. stamineus E. Desv. or as B. cebadilla Steud. Recent morphological 
evidence supports its recognition as a variety of B. catharticus. Detailed illustrations are included 

along with a qgnatee lee description, synonym list, geographical distribution, and representa- 
tive specimens. 


Key Worps: Bromus, Ceratochloa, Poaceae, Bromeae, South and North America 


RESUMEN 


Se propone la nueva combinacion Bromus catharticus var. elata (E. Desv.) Planchuelo. Este taxon fue 
previamente tratado indistintamente como B. stamineus E. Desv. 0 B. cebadilla Steud. Evidencias 
mor os recientes vette su ubicacion como una waned ad ae B. anti El trabajo se 


PAA 


a eres aes distribucion 


senate y especimenes representativos. is 

During the course of studying materials of Bromus deposited in different her- 
baria, several specimens from AHUC (Holmgren et al. 1990) were sent by the 
curator for identification. Some of the specimens agree with the morphological 
features of a native species of Bromus from South America which was identi- 
fied by Camara Hernandez (1978), Matthei (1986), Nicora and Rugolo de Agrasar 
(1987), and Zuloaga et al. (1994), as B. stamineus E. Desv., and by Gutiérrez and 
Pensiero (1998), Planchuelo and Peterson (2000), and Pavlick et al. (2003) as B. 
cebadilla Steud. The native area of distribution of this taxon lies primarily in 
the Southern Andes of the Patagonia region of Argentina and Chile (Gutiérrez 
& Pensiero 1998). In Chile it grows along the Andes and coastal plains (Matthei 
1986), and on Robinson Crusoe Island in the Juan Fernandez archipelago (Baeza 
et al. 2002). In Argentina it grows in western Patagonia (Camara Hernandez 
1978) and on the island of Tierra del Fuego (Moore 1983). 

This grass was introduced many years ago in Central California for experi- 
mental purposes (Hall 1955) and it is now naturalized in North America as re- 
ported in Munz and Keck (1959); Wilken and Painter (1993), Kartesz (1994), 
Pavlick (1995) and Pavlick et al. (2003). Herbarium specimen observations pro- 


SIDA 22(1): 555 — 560. 2006 


556 BRIT.ORG/SIDA 22(1) 


vide testimony that it isa common garden and orchard weed, growing in dis- 
turbed soils in Central and Northern coastal areas of California in the vicinity 
of San Francisco. Also, there are records of it in northern Oregon and southern 
Washington (Pavlick 1995). It has also become naturalized in New Zealand 
(Forde & Edgar 1995), under the names B. stamineus and B. valdivianus, having 
been introduced as part of a program of seed sample trials to screen different 
grasses for their potential forage value. 

Phenetic analyses, based on 16 morphological characters, of types and rep- 
resentative specimens of different taxa of sect. Ceratochloa (including B. ceba- 
dilla) (Planchuelo 1991), indicated that B. cebadilla and B. catharticus should 
be considered conspecific varieties, as suggested by Peterson and Planchuelo 
(1998). This variability within this group of grasses has led to the description 
of numerous taxa at specilic and varietal levels based on size, number of nerves, 


pubescence of the glumes and lemmas, and the length of the lemma awns. Peter- 
son and Planchuelo (1998) clarified the nomenclatural ambiguity of some of 
the names by accepting the new combination B. catharticus var. rupestris, leav- 
ing the status of other taxa for future investigation. Massa et al. (1997, 2001, 
2004)—on the basis of their molecular and morphological studies of 30 


i 


germplasm accessions from Patagonia—recommended treating octoploid mem- 
bers of the B. catharticus complex as a distinct species, B. coloratus, and the 
hexaploid members as B. catharticus. Within B. catharticus, they recognized two 
subspecies, subsp. catharticus and subsp. stamineus. | compared the morphol- 
ogy and verilied the application of the relevant names by examining type ma- 
terials for members of the Bromus catharticus complex. Recognition of B. 
stamineus and B. cebadilla at the varietal level seems warranted. At this rank, 
the epithet elata, based on Bromus unioloides var. elata E. Desv., a taxonomic 
synonym, has priority. A detailed description, synonyms, representative speci- 
mens, geographical distribution and illustrations of Bromus catharticus var. 
elata (E. Desv) Planchuelo are provided below. 


Browns cathartieus var. elata (E. Desv.) Planchuelo, comb. nov. (Fig. 1). Basiony™: 
svar. clatd E. Desv. in Gay, Fl. Chil. 6:438. 1854. Bromus unioloides ft. elatus CE. 

Desv.) Allen & Thell. ex Kloos, Ned. Kruidk. Archief 1917:164. 1918. Type: CHILE: Santiago, C. 

Gay s.n. (LECTOTYPE designated here: Kk! ISOLECTOTYPE: P; photo ACOR!, CONC), SGO). SYNTYPES 
intext: CHILE: Santiago, C. Gay s.n.; Andes de Santa Rosa, FE. Poe ppig s.n; Concepcion, J.8.C.D. 


d’Urville s.n. 


Bromus cebadilla Steud., Syn. Pl. Glumac. 1:321. Apr 1854. Type PROTOLOGUE: CHILE: Rancagua, 

828, Bertero 117 (LECTOTYPE designated here?: P. fragm. ex P. BAA | US-805524!; photo SGO, 
CONC, ISOLECTOTYPE: MO!). SYNTYPES in text: CHII na Ins. Juan Fernand., C.G. Bertero LI7, 118, 
Sol, 1411 (Bertero . sh paw pe of Bri us E. Desv.) 

Bromus stamineus E. Desy. in Gay FL. Chil. 6: 440, 1854. Ceratochloa staminea (E. Desv.) Stace, 
Watsonia 18:413. a esate catharticus subsp. stamineus (E. Desv.) Massa., Canad. J. Bot. 

36-144. 2004. TYPE PROTOLOGUE: CHILE: Rancagua, 1829, C. G. Bertero HI7 (HOLOTYPE: P, 

fragm. ex P US 865470! IsoTyPE: MO! photo, CONC, SGO). 


@) 


i) 


557 


t 


PLANCHUELO, BROMUS CATHARTICUS COMPLEX 


la 


Wr) 


Fig. 1.B 


JIINIAU 
\ 


elata.A. Habit.B, C, D. Spikelets. E. Floret. A, B, E, /lin 73 (CORD); C, Penal 


D, Howell 29056 (AHUC). 


BRIT.ORG/SIDA 22(1) 


Bromus valdivianus Phil., Linnaea 29:102. 1858. Ceratochloa valdiviana (Phil.) Holub., Folia Geobot. 
Phytotax. 8:71. 1973. TYPE: CHILE. Prov. Valdivia: R.A. P hilippis.n. HOLOTYPE: SGO-PHIL-463). 


Usually perennial, occasionally annual, caespitose. Culms 40-100 cm tall, base 
of the plant with some sheaths from previous seasons. Sheaths glabrous or 
slightly pubescent open on the upper quarter part. Ligule membranous, gla- 
brous, 2-4 mm long, apex dentate. Auricles absent. Blade 10-30 cm « 4-7 mm, 
flat, glabrous or slightly pubescent on the upper side. Panicles erect, semi-con- 
tracted, 10-20 cm long, with 15-30 or more spikelets, pedicels 2-10 cm long. 
Spikelets 15-25 mm long, with 4-7-florets, florets imbricate, laterally com- 
pressed. Glumes glabrous, the first glume narrowly ovate-lanceolate, 6-8 mm 
long, 5(-7)-nerved, occasionally only 3 nerves conspicuous, 2 additional nerves 
evident at the base, the second glume ovate-lanceolate, 7-10 mm long, 7(-9)- 
nerved. Lemmas ovate-lanceolate, glabrous, usually smooth, occasionally sca- 
brous on all parts, 10-12 mm long, 7-9-nerved, apex bi-denticulate, the awn 
sub-apical, straight 5-10 mm long. Paleas 10-12 mm long, strongly keeled, with 
ciliate nerves and adherent to the caryopsis. Anthers 3-4.5 mm long. Caryopsis 
7-8 mm long, with a deep, narrow furrow. 

Distribution and habitat.—Native to South America, B.catharticus var. elata 
grows along the Cordillera de Los Andes from Peru to western Patagonia of 
Argentina and Chile, and on the islands of Tierra del Fuego and Robinson 
Crusoe. It isadventive in North America where it occurs in Central and North- 
ern coastal areas of California. Pavlick (1995) reported that it also grows in north- 
ern Oregon and southern Washington but no records substantiating his report 
are known (Barkworth, pers comm. 2006). Reported as naturalized in New 
Zealand (Forde & Edgar 1995). Grows in disturbed soil, waste places. 

Common names.—In Argentina “cebadilla,” in Chile: “lanco,” “Hanco,” “ce- 
badilla,” “pasto del perro” (Matthei 1986) and in USA “Chilean brome” (Pavlick 
1995) 


Representative specimens see ARGENTINA. Chubut: Dpto. Cushamen: Cholila, 15 Jan L901, II lins 


192(CORD). Dpto. Futaleufu: Futalaufquen, 9 Jan 1948, Sor iano 2868 (BAA), 3 Feb 1955, Burkart 
1978] (SI, US); 5 Feb 1955, oe (SI, US). Dpto. Languineo: Region ee rio Corcovado, 20 Dec 
1901, Illin 28 (CORD); 71° W 43° S, 4-6 Mar 1901, [lin 73 (CORD). ee : . Lacar: Parque Na- 


cional Nahuel Huapi, Lago Traful, 7 Nov 1949, Boelcke etal. 3646(BAA); San ee de los Andes, Dec 
1938, Parodi 13212 (BAA). Rio Negro: Dpto. Bariloche: Lago Nahuel Huapi, 900 msm, 8 Feb 1934, 
Parodi 11748 (BAA). Dpto. Bariloche: Bariloche, Parodi 155691/2 (BAA). Cerro Catedral, Dec 1961, 
Ellenberg 1059 (BAA). CHILE. II Region de Antofagasta: Antofagasta, Quebrada Cerisso, road to air- 
port S of Antofagasta, 27 Feb 1939. Beetle 26188 (MO, US). HIT Region de Atacama: Atacama, 27 Fe 
1939. Beetle 26188 (MO). 1V Region de Coquimbo: Dpto. Illapel, Choapa, desvio Pola hasta tunel, 1200- 
1350 msm, 12 Oct 1945. Biese 2128 (LIL). V Region de Valparaiso: Valparaiso, An Wegen, 1895, Buchtien 
s.n.(US); Zapallar, | Feb 1920, Holway et al. 308 (US). VI Region del Maule: Maule, Arroyo 96089 (CONC, 
MO). Constitucion, Oct 1891 Philippi s.n. (US). VIL Region del Bio Bio: Concepcion: 29 Oct 1919, Hol way 
et al. 150 (US); Oct 1925. Claude-Joseph 179 (US). 1X Region de la Araucania: Cajon, Dec 1942, Claude- 
Joseph 5812(US); X Region de los Lagos: Valdivia, 1888. Philippi s.n.CUS). PERU. Junin: Tarma, 11° 50'S 
75° 56’ W, 3780 msm, 7 Jan 1983, Smith 2980 (MO). La Libertad: Trujillo, Cerro Cabezon, 500 msm, 4 


ion 


PLANCHUELO, BROMUS CATHARTICUS COMPLEX 559 


Nov 1983, Sagastegui & ee 005 (MO). UNITED STATES OF AMERICA: CALIFORNIA: Alameda 
. Gill Tract Albany, 5 Junl944, Stebbins Jr 436-1 (AHUC),; Berkeley, 29 Apr 1916, Smith s.n, AHUC 
2029 AHO 14 May 1942, Beetle 3267 (AHUC); 27 May 1942, Harlan 2745 (AHUC); U.C. Campus, 
‘ford st. opposite University Ave., 3 Jun 1924, Buckley 14 (AHUC); in front of ae near Oxford, 4 
en Kennedy s.n, AHUC 2027, 2028(AHUC). Fresno Co: Kings River Canyon near Copper Creek, 
5000 ft elevation, 2 Aug 1958, Howell 34232 rs Humboldt Co.: Fortuna, Fair ground’s experi- 
ieee area, 19 Aug 1947, Murphy s.n, AHUC 26298 (AHUC). Mendocino Co.: Van Damme State Park, 


endocino and ca. | mi up Little River from Van Damme Beach, along Fern Canyon trail, 14 Jul 1988, 
ee see AHUC). Marin Co.: Tiburon Blvd., 150 yards E of Blackfield Dr, 26 Aug 1961, Penalosa 
2220 (AHUC). Napa Co.: Oakville, Elev. ca. 160 ft, 20 May 1951, Raven 2836 (AHUC); Calistoga, alti- 
tude ca. 300 ft, 5 May1957, ae 10788 (AHUC). Sacramento Co.: Brown's Ranch, Elk Grove, 25 Apr 
1947, Cakins.n., DAV 10032 (AHUC ). San Luis Obispo Co.: Adelaide Road, 4.7 mi W of Paso Robles, 
1700 ft, 21 May 1960, meal nn 5970(AHUC). San Francisco Co.: San Francisco, Jackson Street, 2 
Jun 1953, Howell 29056 (AHUC). Sonoma Co.: Bluffs above the sea, Sonoma Coast State Park, along 
State Hwy. 1, 0.9 mi N of Salmon Creek, 11 Jul 1957, Crampton 4326 (AHUC). Yolo Co.: UC Davis 
agronomy experimental area, plant introduction nursery, May 1950, Crampton s.n. AHUC 042728 
(AHUC). 


Planchuelo and Peterson (2000) recognized eight species in the South Ameri- 
can members of sect. Ceratochlod. Massa et al. (2004) proposed recognizing only 
two species, B. catharticus and B. coloratus, with two subspecies within B. 
catharticus. In this paper, one of the species recognized by Planchuelo and Peter- 
son (2000) as B. cebadilla is reduced to a variety. The status of the other species 
recognized in the mentioned paper merits further study, 


ACKNOWLEDGMENTS 


The author thanks Mary Barkworth for the loan of specimens and her construc- 
tive comments on the manuscript, Victor Finot for sending photographs and 
information of type materials and to the curators of the listed herbarium for 
sending specimens on loan. Kanchi Gandhi provided useful comments on the 
nomenclature. 


REFERENCES 


Baeza C.M., T.F Stuessy, and C. Marticorena. 2002. Notes on the Poaceae of the Robinson 
Crusoe (Juan Fernandez) Islands, Chile. Brittonia 54:154-163. 

CAMARA HernAnoez, J. 1978.Bromus.In:M.N. Correa, ed. Flora Patagonica. Coleccion Cientifica 
INTA 8 (3):77-93. Buenos Aires. 

Forbe, M.B. and E. Edaar. 1995. Checklist of pooid grasses naturalised in New Zealand. 3. 
Tribes Bromeae and Brachypodieae. New Zealand J. Bot. 33:35-42. 

Gutierrez, H.F.and J.F Pensiero. 1998. Sinopsis de las especies argentinas del género Bromus 
(Poaceae). Darwiniana 35:75-114 

Hatt, B.M. 1955. Genetic analysis of interspecific hybrids in the genus Bromus, section 
Ceratochloa. Genetics 40:1 75-192. 

Houmeren, P.K., N.H. Houmeren, and L. Barnett. 1990. Index herbariorum. Part I:The herbaria of 
the world. 8th ed. New York Botanical Garden, Bronx, NY. 


= 


560 BRIT.ORG/SIDA 22(1) 


Kartesz,J.1.1994.A synonymized checklist of the vascular flora of the United States,Canada 
and Greenland Volume 1 Checklist. Timber Press, Portland, OR. 

Massa, A.N., K.B Jensen, S.R Larson, and DJ. Hott. 2004. Morphological variation in Bromus 
sect. Ceratochloa germplasm of Patagonia. Canad. J. Bot. 82:136-144. 

Massa, A.N., S.R. Larson, K.B. Jensen, and D.J. Hote. 2001. AFLP variation in Bromus Section 
Ceratochloa Germplasm of Patagonia. Crop Sci. 41:1609-1616. 

Massa, A.N., A.H. Zappe, R. GNDULLO, H. Acuna, and I. Secuet. 1997. Collecting Bromus L.in the 
Patagonian Andes. Plant Genet. Resour. Newsl. 110:1-4. 

Mattel, O. 1986. El genero Bromus L. (Poaceae) en Chile. Gayana 43:47-110. 

Moore, D.M.1983.Flora of Tierra del Fuego. A.Nelson-Missouri Botanical Garden, England- 


ts 


Munz, RA.and D.D. Keck. 1959.A California flora, Supplement, 1968; Combine edition 1973. 
University of California Press, Berkeley. 

Nicora, E.G. and Z.E. RUGoLo pe Acrasar. 1987.Los géneros de Gramineas de América Aus- 
tral. Hemisferio Sur. 

Pavuck, L. 1995. Bromus L. of North America. Royal British Columbia Museum, British Co- 
lumbia, Canada. 

Pavuick, L.E., AM. PLANCHUELO, PM. Peterson, and R.J. Sorenc. 2003. Bromus. In: RJ. Soreng, P.M. 
Peterson, G. Davidse, E.J. Judziewicz, FO. Zuloaga, T.S. Filgueiras and O. Morrone, O. eds. 
Catalogue of New World grasses (Poaceae): IV. Subfamily ior Contr. US Natl. Herb. 

48:1—730, Smithsonian Institution. Washington, DC. Pp. 154-189. 

Peterson, P. and A.M. PLancHueLo. 1998. Bromus catharticus in ets America (Poaceae: 
Bromeae). Novon 8: . 

PrancHuelo, A.M. 1991, Estudios sobre el complejo Bromus catharticus (Poaceae), |. 
Evaluacion estadistica de los caracteres taxOnomicos. Kurtziana 21:243-257. 

PLANCHUELO, A.M.and P.M. Peterson. 2000. The species of Bromus (Poaceae: Bromeae) in South 
America. In:W.L.J. Surrey and J. Everett, eds. Grasses systematics and evolution. CSIRO 
Publishing, Australia. Pp. 89-101. 

Witken D.H.and E.L. Painrer. 1993. Bromus.n:J.C. Hickman, ed. Jepson manual: higher plants 
of California. Univ. California Press, Berkeley. 

ZULOAGA, F.O., E.G. Nicora, Z.E. Rucoto pe AGrasar, O. Morrone, J. Pensiero, and A.M. CiALDELLA. 
1994. Catalogo de la familia Poaceae en la Republica Argentina. Monogr. Syst. Bot. Mis- 
souri Bot. Gard. 47:1-178 


CHROMOSOME NUMBERS FOR WESTERN AND ARCTIC 
NORTH AMERICAN SPECIES OF 
ANTENNARIA (ASTERACEAE: GNAPHALIEAE) 


Jerry G. Chmielewski 


ail 


Department of Biology 
Slippery Rock University 


Slippery Rock, Pennsylvania, U.S.A. 16057 


ABSTRACT 


Chromosome numbers were determined from somatic material for 73 individuals of Antennaria 
Gaertn. (Asteraceae: Gnaphalieae) representing 10 species from western and arctic ee America. 
These counts sup poe ane ee the ty eas more meen prev ate 


number d | though 


they s supplement our know ibdee relative to chromosome num mbar distributions for these species. 
RESUMEN 


Se contaron los numeros cromosomaticos en materi: . somatico Mee rhe) untae de Antennaria 
Je Nc América 


Gaertn. (Asteraceae: G 


ae: 


Estos recuentos apoyan, ; repre edie an ali 
la especie respectiva y como tales son presenti gaae sin ningun comentario, nan complen — 
nuestro conocimiento relativo a las distribuciones de nimeros cromosomaticos de estas especies. 


The purpose of this paper is to supplement the chromosome literature for North 
American Antennaria with previously unpublished data. The methodology 
employed for chromosome number determination was previously described 
(Chinnappa & Chmielewski 1987). Voucher specimens remain in the posses- 
sion of the author and duplicates are deposited in SLRO. The author collected 
all specimens and staminate specimens are designated with an asterisk (*). 


Asteraceae 


Antennaria densifolia Porsild. 2n = 28. YK: Dempster Hwy., Engineer Creek, ki- 

lometer post 178, 65° 14.243'N and 138° 17.714 W, Ch3138, Ch3139*; N of Tomb- 

stone Campground, kilometer post 201, 65° 23.900' N and 138° 16.411 W, Ch3140, 

Ch3141*, approx. kilometer post 213, 65°28'100' N and 138°12.312' W, Ch3142, 
ico 


Antennaria friesiana (Trautv.) Ekman ssp. alaskana (Malte) Hultén. 2n = 28. AK: 
Steese Hwy., vicinity of milepost 108, Eagle Summit, 65° 29.515' N and 145° 
23.314 W, Ch3098, Ch3099*, 65° 30.367' N and 145° 22.704' W, Ch3100, Ch3101*, 
Dalton Hwy., Finger Mtn., 66° 21.465' N and 150° 27.694 W, Ch3108, Ch3109*, 


SIDA 22(1): 561 — 563. 2006 


562 BRIT.ORG/SIDA 22(1) 


Chandler's Shelf, approx. 63 mi N of Coldfoot, 68° 03.026 N and 149° 37.431 W, 
Ch3115, Ch3116*; Atigan Pass, 68° 08.225'N and 149° 26.654' W, Ch3119, Ch3120*. 
Antennaria friesiana (Trauty.) Ekman ssp. friesiana. 2n = 56. AK: Dalton Hwy, 
vicinity of Toolik Lake, 608° 38.411’ N and 149° 30.815’ W, Ch3 118. YK: Top of the 
World Hwy., Boundary rest area, 64° 04.48' N and 140° 58.598' W. Ch3132. 
Antennaria howellii Greene. 2n = 84. AB: Yellowhead Hwy., W of Vermillion, 
53° 20.188' N and 110° 57.004 W, Ch3065; BC: BC 2, Eof Pink Mtn., milepost 140, 
57° O1.454' N and 122° 26.738' W, Ch3068; E of the Yukon border, 59° 57.818' N 
and 127° 29.890' W, Ch3073. MB: Hwy 16, E of Neepawa, 50° 13.546' N and 99° 
O1.443' W, Ch3053. ON: Trans Canada Hwy., W of Spanish River, 46° 12.289'N 
and 82° 23.832' W, Ch3048; W of White River but E of Marathon, 48° 43.143'N 
and 85° 42.543' W, Ch3049; Kakabeka Falls, 48° 23.762' N and 89° 37.925' W, 
Ch3050, Ch3051. SK: Yellowhead Hwy., E of Sheho, 51° 34.483'N and 103° 11.392! 
W, Ch3056. 


Antennaria microphylla Rydb. 2n = 28. AB: Yellowhead Hwy, W of Vermillion, 
53° 20.188) N and 110° 57.004 W, Ch3063*, Ch3064; SK: Yellowhead Hwy., W of 
Langham, 50° 57.243' N and 102° 03.873 W, Ch3055, E of Sheho, 51° 34.483'N 
and 103° 11.392' W, Ch3058*, Ch3059*: W of North Battleford, 53° 03.284' N and 
109° 07.845' W, Ch3061*. 


Antennaria monocephala DC. 2n = 28. AK: Richardson Hwy., Thompson Pass, 
61° 07.899'N and 145° 44.158' W, Ch3079, Ch3080*; Denali Hwy. rest area approx. 
78 mi E of Cantwell, Ch3090, Ch3091*. 

Antennaria pallida E. Nelson. 2n = 56. AK: AK 3, borough of Mirror Lake, 63° 
20.600' N and 149° 05.136' W, Ch3085; Denali Hwy, approx. 21 mi E of Cantwell, 
63° 20.618'N and 148° 17.304' W, Ch3086; approx. 22 mi E of Cantwell, 63° 20.002’ 
N and 148° 16. 166' W, Ch3088; Denali Hwy., approx. +9 mi E of Cantwell, 63° 
10.201 N and 147° 32.652' W, Ch3089, Denali Hwy., rest area approx. 78 mi E of 
Cantwell, Ch3092; Taylor Hwy., approx. 22 mi NE of Teslin Junction, 63° 36.781) 
N and 142° 21. 103' W, Ch3130; YK: South Canol Road, approx. milepost 170, 61° 
41.794' N and 133° 04.206' W, Ch3147. 


Antennaria parvifolia Nutt. 2n = 112. AB: Yellowhead Hwy., W of Vermillion, 
53° 20.188' N and 110° 57.004’ W, Ch3062. SK: Yellowhead Hwy., W of Langham, 
50° 57.243' N and 102° 03.873 W, Ch3054; E of Sheho, 51-34-483°N and 103° 
11.392' W, Ch3057: W of North Battleford, 53° 03.284' N and 109° 07.845' W, 
Ch3060. 


Antennaria pulcherrima (Hook.) Greene. 2n = 56. AK: Dalton | lwy., Coldtfoot, 67° 
15.047' N and 150° 10.580' W, Ch3110, Ch3111*. BC: Alaska Hwy., Muncho Lake, 
59° 09.577' N and 125° 53.319’ W, Ch3070. 


Antennaria rosea Greene. 2n = 56. AB: AB 43, W of Grande Prairie, 55° 10.240'N 


CHMIELEWSKI, CHROMOSOME NUMBERS OF ANTENNARIA 563 


and 119° 16.162' W, Ch3066. AK: Hwy 1, W of Glenallan, 62° 05.729' N and 146° 
14.938' W, Ch3082; Seward Hwy., Turnagin Arm, S of Anchorage, 61° 00.387'N 
and 149° 41.577' W, Ch3083; Fairbanks, intersection of Boat St. and Sportsman 
Way, 64° 50.228' N and 147° 49.445’ W, Ch3096; Elliott Hwy,, S of Livengood, 
65° 09.350' N and 147° 53.893' W, Ch3104; Dalton Hwy., Yukon River, 65° 52.764' 
Nand 149° 42.890' W, Ch3105: N of Yukon River, 65° 57.88' N and 149° 58.931' W, 
Ch3106; Richardson Hwy., North Pole, 64° 45.031 N and 147° 19.524' W, Ch3124; 
Taylor Spur to Boundary, 64° 04.684' N and 141° 06.512' W, Ch3131. BC: Alaska 
Hwy., Muncho Lake, 59° 09.577’ N and 125° 53.319' W, Ch3069; N of Fireside, 59° 
41.772' N and 127° 12.992' W, Ch3071. E of the Yukon border, 59° 57.818' N and 
127° 29.890 W, Ch3072. YK: Alaska Hwy., Watson Lake, 60° 01.172' N and 128° 
34.772' W, Ch3074, Ch3075; N of Dawson Peaks, 60° 26.671' N and 133° 36.476 
W, Ch3076, Ch3077, Top of the World Hwy., W of 60 Mile Rd., W of kp 86, E of 
Boundary, 64° 06.929 N and 140° 43.409' W, Ch3133; Dawson Bonanza Creek, 
gold panning site 6, Ch3134; Dawson, dome overview, Ch3135; South Canol Road, 
Quiet Lake, 60° 59.475' N and 133° 01.678' W, Ch3148, Whitehorse, Miles Can- 
yon, 60° 39.681' N and 135° 01.750' W, Ch3151, Ch3152. 


ACKNOWLEDGMENTS 
The Department of Biology, Slippery Rock University is thanked for their par- 
tial financial support of fieldwork. Dale Johnson and Guy Nesom are thanked 
for their helpful suggestions in their reviews of the manuscript. 
REFERENCES 
Cuinnapea, C.C.and J.G. CHMIELEWwski. 1987. Documented plant chromosome numbers 1987: 
1. Miscellaneous counts from western North America. Sida 12:409-417. 


564 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


LONDA SCHIFBINGER and CLAUDIA SWAN (eds). 2005. Colonial Botany: Science, Commerce, 
and Politics in the Early Modern World. (ISBN 0812238273, hbk.). Univer- 
sity of Pennsylvania Press, Philadelphia 19104-4011, U.S.A.. (Orders: Hop- 
kins Fulfillment Services, PO. Box 50370, Baltimore, MD 21211-4370, US.A,, 
Tel: 410-516-6956; Fax 410-516-6998. e-mail: hfscustserv@mail press. jhuedu) 
[Price not given.] 346 pp. b/w illustrations, locm Xx 25cm. 


ploitation of the world” as the great powers, Spain 


This book might have been titled the “botanica 
and Holland especially, sent off explorers as much for botanical resources as for the conquest of land. 


Their quest was, of course, for the spices and condiments that they were buying from the Far East 


and for the materi dica that had long ae an ee partof botanical pursuit. Botanical knowl- 


edge of the‘ crew areas was limited; tl lificati f plants was imperfect, faulty, and often mis- 


ing: the understanding of biogeography Ww uate with misconceptions. The expedi were 


leadi 
economically and politically motivated with the result of some extreme inhumane ae ieee 


as the atrocities in Banda), anc ee of gee and arrogance the indigenous people’s use ol 
plants for oe purposes was often ignorec 
xteen chapters of this book by as many historians and geographers present particulars 


of ee exploration, aspects of colonial history seldom described. Along the way, one may be- 


come reacquainted with such botanist-physician explorers as Charles-Marie de La Condamine, Jo- 
seph de Jussieu, Carlos Clusius and meet fascinating pier like Nicolas Monardes, who never 
left Spain, and Daniel Gottlieb Messerschmidt. Facts such as the cultivation of African rice (Oryza 
glaberrima) mca into oe and North Carolina by slaves give a new outlook on agriculture and 


life in the s 


For this reviewer more ehomels identifications would have been welcomed, though some au- 
tl | For instance, what was the “tea” that José Mutis pppoe asa sub- 


re “bastard nutmeg” that Pierre Poivre brought from the Philippines 


stitute for Asian tea? Ww hat wast 
to cultivate in France? Answers to these and other such questions woulc 


tensive citations of original and secondary sources for each chapter in the appended Notes. 


surely be found in the ex- 


ao ecilors have ee an elas cera ye essa ye bee more ome fae ex- 
lai ann k arees 


franied TCU Library), Botanical Receacl Institute of Texas, Fort Worth, TX 76102- 4066, U.S.A. 


SIDA 22(1): 564. 2006 


ROOFSHOOT ANATOMY AND POSTHARVEST VEGETATIVE 
CLONAL DEVELOPMENT IN LOPHOPHORA WILLIAMSII 
(CACTACEAE: CACTEAE): IMPLICATIONS FOR CONSERVATION 


Martin Terry James D.Mauseth 
Department of Biology Section of Integrative Biolo 
Sul Ross State University College SiNEA iG Sciences 
pine, Texas 79832, U.S.A. The University of Tex 


Austin, Texas 78712, U.S.A. 


ABSTRACT 
Over the last four decades, the size and density of populations of Lophophora williamsti (peyote) 
have diminished markedly in large areas of South Texas where licensed peyote distributors harvest 
the cactus 


—— 


for ceremonial use by the Native American Church. Part of the problem lies in the fact 


that some harvesters are cutting plants too hou on ae subi ranean stem or taproot. That practice 


precludes the regeneration of new f the decapitated plants. 
To address this problem, we describe the anatomical distinctions between subterranean stem and 
root in L. williamsii as follows: The s EG edisinguelie Pe cortical bundles running 


through the parenchyma, in contrast tex, sists Beale parenchyma wamnout 


cortical bundles. The pith at the center of tl l 


distinguished from the dilatated metaxylem (with masses of dark-staining metaxylem tracheary 
seat cecum the center of the root. With these new anatomical tools, it is now possible to set 


upt s, first in the greenhouse and then in the field, to generate prectica! biomet- 
icd | i | a| ptl tv hi nN the peyote har ves ut the plants without 


vhic 
significantly reducing the survival rate noi the rootstocks left in the eal ce harvest. 


RESUMEN 


Durante las ultimas cuatro décadas, el tamano y eBSIdaG de las poplaciones de Lophophora 
are a \} J; ‘ oN We 


| de Texas donde se 
dan permisos a distribuidores de peyote para secclecs especimenes de uso en las ceremonias 
religiosas de la Native American Church. Parte del problema estriba en el hecho de que algunos 
colectores estan cortando plantas muy abajo, o en el tallo subterraneo o en la raiz pivotante. Esta 
practica impide la regeneracion de nuevos tallos y finalmente ocasiona la muerte de las plantas 
decapitadas. Para abordar este problema, nosotros describimos las caracteristicas anatomicas que 
distinguen el tallo subterraneo y la raiz de L. williamsii como sigue: El tejido cortical del tallo se 
distingue por la presencia de haces corticales que se distribuyen por el parénquima, atributo que 


contrasta con el tejido cortical de la raiz, el cual consiste en puro parénquima sin haces corticales. En 
| lel tallo la médulaes } parénq il ),y se reconoce claramente del metaxilema 


lilatad hes de el les del metaxilema tenidos de oscuro) que ocupa el centro de 


r 
la raiz. En base a estas caracteristicas anatomicas sera posible hacer experimentos de corte, primero 


en invernadero y 1 n la finalidad de generar datos que permitan determinar la 
profundidad maxima a la cual los a me de peyote pueden cortar las plantas sin reducir 
significativamente la supervivencia de los individuos que quedan en el suelo después de la colecta. 


SIDA 22(1): 565 - 592. 2006 


BRIT.ORG/SIDA 22(1) 


INTRODUCTION 

Peyote harvest as a conservation problem 

Lophophora williamsii(Lem.ex Salm-Dyck)J.M. Coult. 1894, commonly called 
peyote, is a small, napiform, spineless cactus native to the Chihuahuan Desert 
and the Tamaulipecan Thornscrub of northeastern Mexico and adjacent Texas. 
While the mescaline-containing stems are ingested in various forms for their 
psychotropic effects, this behavior was generally proscribed by federal drug 
legislation in 1970. However, federal law provides protection for the use of peyote 
for bona fide religious ceremonial purposes by members of the Native Ameri- 
can Church. The supply of peyote for such purposes is regulated by the Drug 
Enforcement Administration and the Texas Department of Public Safety. The 
regulated commerce in peyote begins with the harvest of peyote from wild popu- 
lations by licensed peyote distributors or their agents. Commercial quantities 
of peyote occur in the US. only in Starr, Zapata, Webb and Jim Hogg Counties 
in South Texas, within 70 km of the Rio Grande, and all four currently licensed 
peyote distributors are based where peyote grows in those counties. Historically 
the peyote distributors have gained access to harvestable populations of peyote 


— 


through peyote-specilic lease ag ents with private landowners. Most of the 
actual harvesting of peyote is done by contract laborers who are paid in accor- 
dance with the number and size of freshly cut “buttons” (tops of stems) of peyote 


that they deliver to the licensed distributors. 

There has been a decrease in the number, size, extent and density of peyote 
populations in South Texas (Anderson 1995; Moreno 2005) over the past four 
decades. Much of this reduction in peyote numbers can be attributed to habitat 
destruction associated with urban development and agricultural practices such 
as rootplowing of native brush, and some adverse effects on population num- 
bers may have been due to illicit harvesting. Such phenomena are difficult to 
quantify and virtually impossible to prevent. One factor that is quantifiable in 
the decline of peyote in South Texas is the regulated commercial harvest of 
icensed distributors for ceremonial use by the Native American 


— 


peyote by the 
Church. Approximately two million peyote buttons per year have been har- 
vested by these distributors over the last two decades (Texas Department of 
Public Safety, unpublished data). And the technique used in the harvest of those 
plants could logically have a substantial impact on the observed decline—or 
the potential recovery—of peyote populations in the Tamaulipecan Thornscrub 
of South Texas. 

The proper technique for harvesting peyote is that the crown (ie., the aerial, 
photosynthetic portion of the stem) of the peyote cactus is cut off at or imme- 
diately below its base, and the subterranean portion of the plant, including all 
or most of the subterranean portion of the stem, is left in the ground to regen- 
erate one or more new crowns. Such harvesting of the commercially valuable 
crown of the cactus may be accomplished by cutting through the plant trans- 


versely at the level of the surface of the ground, at or near the interface of the 
green crown and the brown subterranean portion of the stem, using a machete, 
a cutting tool with a broad flat blade and a handle about 60 cm long (such asa 
hand edger), or virtually any kind of knife (M. Terry, pers. obs., Fig. la-c). 

When proper harvesting technique is adhered to, the decapitated subter- 
ranean portion of the stem and the more distal taproot of the cactus (Fig. 2) 
remain intact and in situ, where the viable subterranean stem tissue will nor- 
mally begin to regenerate one or more new crowns by lateral branching from 
axillary (areolar) buds within a few months after loss of the apical meristem 
(Fig. 3). Such regenerated crowns may in turn be sustainably harvested alter 
they reach maturity, some years later (Fig. 4). 


Peyote harvest—degeneration and death of the decapitated plant 

Not every peyote plant responds in the same manner to removal of its apical 
meristem along with the crown at harvest. The simplest response is that de- 
scribed above: regeneration of one or more new crowns by lateral branching 
from (usually subterranean) stem tissue (Fig. 3). But frequently we observe a 
more complex, gradual response, which begins with simple lateral branching 
from the remaining stem of the decapitated plant. The difference is that the new 
stem branches, instead of remaining dependent on the taproot of the original 
plant, put down their own adventitious taproots and eventually become inde- 
pendent plants that detach themselves from the original plant, which—if its crown 
was completely removed at harvest—degenerates and dies in this process. 

The unusual aspect of this second type of response of peyote to removal o 
its apical meristem in the harvesting process, is that the regeneration of new 
stems and the generation of new adventitious taproots from the new stem 
branches proceed in a seamless developmental process until at some point it 
must be recognized that the new, increasingly hoot-root units have 
become independent vegetative clones of the original plant. That is to say, where 
we started witha single decapitated plant undergoing development of new stem 
branches, we end up with what must be recognized as a moribund parent plant 
with viable clonal progeny. 

Another remarkable feature of this process is that as the clonal progeny 
become larger and more nutritionally independent, the stem tissue connection 
between the parent plant and each of the vegetative clones degenerates toa slen- 
der tube consisting mostly of vascular tissue, while the parent plant (which 
has no photosynthetic capabilities of its own) steadily decreases in size and 
density as the nutrients stored in its parenchymal tissues are depleted and uti- 
lized by the growing clonal offspring. In the final phase of the process, the con- 
nection between the parent plant and the now nutritionally independent prog- 
eny disintegrates, and what is left of the decapitated parent plant dies. This 
series of events is depicted in Figures 5-12. All the photos in these figures are of 


mS 


568 BRIT.ORG/SIDA 22(1) 


i.e. at ground level 
g 


Fic. 1a. Proper 


| 
oe J “I 7 


specimens that the first author collected from populations near Rio Grande City, 
Starr County, Texas. 

This natural process of vegetative clone production in response to the ex- 
cision of the apical meristem that occurs when peyote crowns are harvested, is 
remarkably congruent with the folk belief that harvesting peyote results in an 
increase in the number of plants ina population, and that where one plant grew 
before harvesting, several plants may be found after allowing an adequate pe- 
riod of time for the peyote to “grow back.” A necessary condition for this pro- 


—_ 


cess to occur, however, is that the harvesting be done ina manner that does not 
oreclude the production of new stems from the subterranean stem of a plant 


from which the crown has been harvested. What does this mean in terms that 
could serve as a practical guideline for sustainable harvesting? 

The answer must be based on an understanding of the anatomy of the root 
and stem of Lophophora williamsii. We begin with the observation that of all 
the people working with this plant—and here we include peyote distributors, 
members of the Native American Church, cactus hobbyists in countries where 
peyote cultivation is legal, and even botanists—very few appreciate the distinc- 
tion between true root and subterranean stem. Yet this distinction is crucial to 


aLN . = L 4 -— L 1 J IlAl + £. J 1 | {hk | 
Fic 


about 5 cm in diameter, with 8 ue) has been ue from subterranean portion of stem mye: poate in the 
ground). C f 1 pith within the 


J / HP 


vascular cylinder, and yellowish green arene fe it ane) cortex of the stem (between vascular ring and 
epidermis). 


— 


an understanding of how to harvest peyote so that it will “grow back,” because, 
as far as we have been able to determine from observations to date, new stem 
branches will develop only from stem tissue, not from root tissue. Therefore, if 
in harvesting the crown one cuts so deeply below the crown that all or most of 
the subterranean stem tissue is removed along with the crown, then there will 
be no possibility of new stem development, and the stemless root left in the 
ground will simply perish. With that as the operating premise, let us now con- 
sider the anatomy of the root and the shoot of L. williamsii, and how to distin- 
guish between them. 

Ideally we should like to be able to identify the shoot-root transition zone 
in order that at least some portion of the shoot should remain with the root of 
the plant left in the ground after harvest. Unfortunately, the plants normally 
protrude only 1-3 cm above the surface of the ground and have a large subter- 
ranean shoot that tapers gradually until it ends in a taproot. The shoot-root 
transition zone does not occur near the soil level where the seed germinated 


570 BRIT.ORG/SIDA 22(1 


= 


Fic. 1c P } H IInd | £} tod (loft) ch 197 3mmw 


of ue at perimeter o cut surface, indicating inet the cut was parely pelo pase ot crown, in uppermos nee of 


eww higher chloro- 


L he + I f Pavia tare, Tae TT * aL 
J 


phyll content 
p 7 t 


but instead occurs at various depths (higher in smaller plants, lower in larger 
plants). It is not possible to identify the shoot/root transition zone merely by 
examining an intact plant with the naked eye. 

In most seed plants, young shoots and roots can be distinguished from eac 


1 
other because the shoot has a pith and cortex whereas the root lacks both these 
structures (Mauseth 1988). However, several other cacti grow like peyote—having 
a large below-ground shoot that tapers into a large taproot—and the shoot-root 
nature of those structures has not been clarified (Stone-Palmquist & Mauseth 
2002). Early workers on the anatomy of Lophophora (Rouhier 1927; Bravo 1931: 
Janot & Bernier 1933) gave lair to very good anatomical descriptions of root and 
shoot, but even Bravo, whose description of the subterranean stem and root ol 
Lophophora was the best available in its day, admitted that ‘it is most difficult to 
know in which region the stem ends and in which the root beginsl..” (Bravo 1931, 
translated from the Spanish). Because better harvest techniques may aid the sur- 
vival of this species in areas subject to intensive harvesting for human consump- 
tion, as in South Texas, we undertook a histologic study of the anatomy of roots 


TERRY AND MAUSETH, ROOT AND SHOOT ANATOMY OF LOPHOPHORA WILLIAMSII 


Fic. 2. Peyote plant immediately after most of crown has been harvested, showing cut just above base of harvested 
{ fd + \ ht + {b h L 14; H + } ¢£ \ 


4 A a) Yee aL £, Te Pd | | \ 


pable of regenerating new crown(s 


and shoots in L. williamsii to determine if there are reliable criteria for distin- 
guishing the root from the shoot (and particularly the subterranean portion of 
the shoot). We especially looked for criteria that could be used in the field. 
MATERIALS AND METHODS 

Plants were collected by M. Terry (DEA Researchers Registration No. RTO269591) 
from a wild population of L. williamsii near Rio Grande City in Starr County, 
Texas, or, in the case of one specimen (the very large plant), donated for research 
purposes by law enforcement personnel. Specimens examined included two very 
small plants, two plants of intermediate size and one very large plant (Table 1). 


572 BRIT.ORG/SIDA 22(1) 


Taste 1.Dimensions of the three adult plants studied, listed from shortest to longest. All values are 
in nee. Shoots of plants "1212 and "1213 were sampled at ground level and at two below- 
ground levels; shoot of plant #1214 was sampled at only one below-ground level. The root — 


plant was sampled only at the top of the root, where it most resembled a portion of sho 
Sample Plant Aerial Plantdiam. Shootcortex Shoot pith Root cortex- Root pith- 
length height atsoillevel thickness diam. like zone like region 


thickness diam 


Plant 1212,at ground = 135 24 os 20 | 
level 

1212, at -20 mm 13 6 

1212, at -25 mm 9 6 

1212, root, -35 mm 1.5 10 
Plant 1214,at ground = 140 44 78 32 10 
level 

1214, at -35 mm 6 27 

1214, root,-41mm 3.0 2/ 
Plant 1213,at ground ~——-160 40 59 14 6 
level 

1213, at -20 mm 14 6 

1213,at-35 mm 9 8 

1213, root, -45 mm 1.0 3.4 


Plants were dissected with care being taken to obtain samples of material 
that was definitely root (provisionally defined as the region below the upper- 
most point at which a lateral root had emerged), definitely shoot (namely the 
aerial portion of the shoot—specifically known as the crown, sensu Schultes 
(1938)—which has a blue-gray to blue-green epidermis, photosynthetic tissue 
and axillary buds), and definitely hypocotyl (the transition zone between shoot 
and root) samples being obtained by taking numerous samples between obvious 


root and obvious shoot). In all but the smallest, youngest plants, the plant ma- 


terial that was easily recognizable as root was located at least 35 mm below soil 
level (45 mm below in plant “1213; Table 1). Since the nature of the higher sub- 
terranean portions of L. williamsii was in question, samples were taken from 
all plants examined, with the position of each sample being carefully measured 
with the soil level (taken as the level of the base of the crown) as reference. 

As tissue samples were obtained during dissection, they were immediately 
immersed in Navashin’s solution, then aspirated ina vacuum chamber to remove 
air and permit rapid penetration of fixative. Tissues were fixed for 24 hours, de- 
hydrated through mixtures of ethanol and tertiary butanol, then embedded in 
Paraplast Plus. After microtoming, sections were stained with Safranin and Fast 
Green by a procedure designed especially for cacti (Mauseth et al. 1985). 


Fic. 3. Two young peyote crowns (“pups”) regenerating by | | branching f he upper edge of th 
+ £ | L J r+ 1756 4 I y f 4 + ol SS 1 L * r: 1a—1c.Each 


J 


of the new crowns is ca. 1.5 cm in diameter 


RESULTS 

Morpholo 
All plants tapered gradually from an unbranched aerial shoot to a region of 
subterranean shoot, then to hypocotyl, and finally to taproot. The taper was 
uniform in most plants, without any abrupt change in diameter that might in- 
dicate the boundary between shoot and hypocotyl or between hypocotyl and 
root. The two smallest plants that we examined (whose above-ground, photo- 
synthetic crowns were 15 mm in diameter) were only 31mm long (9 mm above 
ground, 22 mm below ground) and 50 mm long (10 mm above ground), so seeds 
must have germinated at or slightly below ground level. However, in the three 
adults we examined, the root/shoot junction was located at least 35 mm below 
the soil level (45 mm below in plant *1213), so plants of Lophophora williamsii 
must have contractile roots pulling the root/shoot junction deeper as the plant 
ages. All above-ground portions of shoot were covered with a blue epidermis: 
all subterranean portions were covered with thin, flaking brown bark. Ribs and 
axillary buds (often called areoles in cacti) were obvious on all above-ground 
portions, and withered areoles were occasionally detected (as much as 17 mm 
below soil level on plant *1213). 

Slender lateral roots 1-3 mm diameter) emerged from taproots, but were 
extremely sparse, with only two or three present on any plant. This could be in 


574 BRIT.ORG/SIDA 22(1) 


Fic. 4. Repeated harvesting of successive from th t. Individual on left t hes ( 
rows) inalcating stem eee an ban its crown n harvested thee times in ie past. individual on right has 


neve examination). 


be | 


part because most lateral roots in this species are ephemeral (emerging in re- 
sponse to moisture, then being shed in conditions of drought), small, and frag- 
ile (M. Terry, pers. obs.). The result is that all but the few largest lateral roots are 
broken off and left in the ground when one uproots the plant from its natural 
growth site in habitat—no matter how carefully one goes about extracting the 
plant. When one grows the plant in loose, friable soil under artificial condi- 
tions, more lateral roots and their finer branches remain intact when the plant 
is depotted. The plants used in this study, however, were recently uprooted from 


taL are ee | la+ | 4 J heat Hr £ H n 


Fic. 5. This i te pl i h 

symmetry of the annular constrictions on the long subterranean stem suggests reduced growth rates in periods of 
winter and/or drought. The crown of this mature plant measures ca. 6 cm in diameter and has eight ribs. The shallow, 
elongated indentation in the side of the crown facing the reader is a scar from tissue sampling for DNA analysis 18 


months previously. 


the gravelly soil of their habitat and this would account for some reduction in 
the number of intact (and therefore observed) lateral roots. 

Lateral roots emerged only from areas that were later shown to be taproot 
or hypocotyl, not from shoot tissue. But one plant examined in the field but not 
dissected and studied here had a root emerging from its side at about the same 
level as a lateral stem branch; that root might have emerged from the hypo- 
cotyl rather than the shoot but that is not known. This isolated field observa- 


576 BRIT.ORG/SIDA 22(1) 


Fic. 6. This is a good example of aplant that wasde- _—Fic. 7. This plant st id f having | iously | 


capitated exactly once, several years ago, judging —_ vested at least twice. The small sizes of the two crowns on lat- 
from the size of the new crown (ca. 5 cm diameter). Ik hesf he original sut ggest tl 
Note that the crown-bearing lateral branch is mark- I I ps 2-4 years 
edly offset from the center of the original subterra- ago. (The larger of tl is about 2.5 cm in diameter 
nean stem seen at the base of the lateral branch. Both are 5-ribbed.) 

That is a consequence of the fact that the lateral 

branch originated from an areole on the side of the 

subterranean stem. 


J 
} L * rary eed 1 L 


tion should be interpreted in light of the possibility that some of what appear 
to be uppermost lateral roots of Lophophora may prove to be adventitious roots 
emerging from subterranean stem tissue. Whether ordinary subterranean stem 


tissue can produce adventitious roots, or whether adventitious roots can be pro- 
duced only by regenerative lateral branches from a plant whose apical mer- 
istem has been removed, is an anatomical issue still to be resolved. 


TERRY AND MAUSETH, ROOT AND SHOOT ANATOMY OF LOPHOPHORA WILLIAMSII 


y , but the tw 
x A 


th iwethia ( + 1° + ht) 


Ff J ~s 


al budding from a previous harvest-associated 


J 1 11 | 
MCEVEIVPCU My latel 


tself decapitated 1 level. The two new crowns, each with its own adventitious tap- 
root, are well on their y I i g ind t of tl | k. Tk igi it Ipl i represented 
by its bark I tay I ling tol from the original sut (also bark-covered), is degen- 
erating, but I tem is still ali ill hed I t fthet | | g 


? 


BRIT.ORG/SIDA 22(1) 


Fic. 9. Two vegetative clones that originated as lateral branches of the stem of a decapitated parent plant, which is 
J ed L Pearl | 1 2 hl +h kh L d he oh a = | kh 


J a / J F 
4 4 Jal + raras 4 | + ba 4 aril 4 i 1 thy 1 L £, | ia 
the inter-adherence of the plants in the photo) t I pl 1 the | | plant, but each of the young 
1 L * f, 4 1 4 ¥ W * j J Leh 1 + te t BF eH | 
1g es - P é J 
th i ibuted to tt ivity of | herbi probably terrestrial snails, which are known to consume L. 


williamsii crown tissue. 


Fig. 10. This pair of “sister” vegetati i in Fig 
ac affactivealy 1 +} 4 . 
as Ue aT ad F Li 
2 Ld 


Several plants had shrunken, withered tubercles on their subterranean 
portions. They were wrinkled and covered with bark just like all other subter- 
ranean portions, but their centers contained living parenchyma cells and an 
apical meristem, apparently the shoot apical meristem of the tubercle. The low- 
est one found on each adult plant was 15 mm below soil level on plant *1212, 30 
mm on plant “1213, and 22 mm on plant #1214. 

Withered tubercles were identifiable on subterranean, bark-covered por- 


580 BRIT.ORG/SIDA 22 


1) 


Fic. 11. Three clonal sister peyote plants—each ana 


tomically and functionally on its way to independence with its 


n +h + + £4h tal Cpe set Dearne lei | tc\ 
P y Plalits). 


tions of two of the adult plants. Tubercles were reliably identifiable only in the 
uppermost 4.0 mm of subterranean shoot; below that, they had withered so 
much that the only visible remnants were peg-like structures 2.0-3.0 mm in 
diameter and 1.0-2.0 mm tall, which appeared to be deteriorated areolar tufts 
of trichomes. Bark was sufficiently wrinkled and rough that some of its irregu- 
larities resembled withered tubercles, making identification of tubercles diffi- 
cult. Subterranean withered tubercles should have been aligned with the rows 
of tubercles on the aerial portions (Fig. 13), but there were only two cases in 


Fic. 12.The amorphous, decaying (right) that is tenuously attached to the subt t f the live cactus 
(left), f bt t f P r | I pi 1k “fr i i 
the past TL 2 L J 1 if, L n <_e +5] he, + ow fully for med, 


with its own taproot, and is fully indenendent The very ae density of the attached mass que that it is in an 
advanced stage of decomposition. 


582 BRIT.ORG/SIDA 22(1) 


Fic. 13. L. willi ii bercle witt le | i lly trichomes. Th ical 7 f hich th | 


7 J J 7 


which a row of withered tubercles could be identified by their areolar tufts (Fig. 
14). We followed rows of tubercles from the aerial portions of shoots down into 
subterranean portions but usually could not find any identifiable withered 
tubercles, apparently the areolar tufts abscise from the plant. 

Structure of the root 

Very young regions of root (less than 0.5 mm in diameter) had an organization 
typical of most dicots. There was an epidermis, a thin cortex only a few cells 
thick, endodermis and vascular tissue consisting of small bundles of primary 
phloem alternating with arms of protoxylem,and metaxylem occupied the very 
center. An important point is that metaxylem contained significant amounts 
of living xylem parenchyma cells, it did not consist entirely of dead tracheary 
elements. This organization was found in only the two smallest, youngest plants, 
in samples taken from closest to root tips. Older portions of roots (2.5 to 30 mm 
in diameter) had altered their organization. They had lost their epidermis and 
cortex, and parenchyma cells in the metaxylem had begun to proliferate. Sec- 


Ss 


TERRY AND MAUSETH, ROOT AND SHOOT ANATOMY OF LOPHOPHORA WILLIAMSII 583 


l 1 tal and ay Pare ) l . £ / . . half af 
Fic. 14. i williamsii t \ Nall of 


photo) 41 £. £4L L a hal€-fnrl \s L tee n Lal 


ang shadow are two a antets colores Se ae Sua ona diagonal curve containing ie tuo tumted areles 


L ¢| TL L € L 4 I ® lafth pe | Tot 


ondary xylem (wood) and phloem were present, but epidermis, cortex and en- 
dodermis had been replaced by a bark consisting of thin flakes of cork. Just 
interior to the bark was a band of parenchyma that appeared to be cortex but 
which was really secondary phloem parenchyma. This cortex-like region was 
about 0.3 to 3.0 mm thick and extended inward from the bark almost to the 
vascular cambium. Thickness was correlated with root size: roots less than 4.0 
mm in diameter had a cortex-like region only about 0.3 to 0.4 mm thick; roots 
about 10-12 mm in diameter had a cortex-like region 1.0 mm thick; and very 
large roots 30 mm in diameter had a cortex-like region 3.0 mm thick. The cor- 
tex-like region was recognizable as secondary phloem only because it had traces 
of collapsed sieve tube members in it. There were no vascular bundles in the 
cortex-like region other than very rare connections with lateral roots, and these 
were oriented vertically rather than radially or tangentially. 


584 BRIT.ORG/SIDA 22(1) 


Metaxylem parenchyma proliferated in roots, producing such abundant 
amounts of parenchyma that the center of the root appeared to have a pith. 
Metaxylem vessel elements were pushed apart (such proliferation in an other- 
wise mature tissue is called dilatation), and parenchyma cells in the innermost, 
first-formed wood also underwent dilatation growth. This pith-like region could 
be identified as dilatated xylem (rather than true pith) by the presence of iso- 
lated vessel elements within it; these were easily visible with a handlens and 
dissecting microscope. In roots about 2.5 to 5.0 mm in diameter, the pith-like 
region was about 0.5 to 1.2 mm in diameter, but it was 3.4 mm in diameter in 
roots 12 mm wide, 10.0 mm in diameter in roots 22 mm wide, and 27 mm wide 
in roots 35 mm wide. Dilatation occurred in both the innermost, first-formed 
secondary xylem as well as the middle regions, but the outer regions of second- 
ary xylem (the outermost L.0-2.0 mm) had ordinary wood organization. 

Root wood consisted of a ray system and an axial system (containing axi- 
ally oriented cells such as vessel elements). Rays were extremely narrow, only 
73sd 2.3 um wide and consisted of large, rounded parenchyma cells. The axial 
system consisted of vessel elements, paratracheal parenchyma in immediate 
contact with the vessels, and wide-band tracheids (WBTs). Wide-band trache- 
ids are an unusual type of cell found in almost all cacti; they are short (range in 
Lophophora: 315 to 525 wm), broad spindle-shaped tracheids with secondary 
walls that are annular or helical (Mauseth et al. 1995; Mauseth 2004). There 
were no fibers in the wood. Just as rays were narrow, so were axial masses (98.4 
sd 54 um wide), and cross sections of root wood appeared to be rather solid 
when viewed with the naked eye or by dissecting microscope. 


=_ 


Structure of the shoot 
Young regions of shoot differed from older regions by still having epidermis 
but lacking secondary xylem, phloem and bark. Epidermis was present on all 
aerial portions of shoots and had a blue-gray color. Hypodermis consisted of 
one layer of parenchyma cells. Both epidermis and hypodermis cells definitely 
did not have thickened walls so the shoot surface was very soft. Shoot cortex 
was always much thicker than the root’s cortex-like region of secondary ph- 
loem. The thinnest cortex in an adult plant was 6 mm (in an old, below-ground 
portion of plant *1214) and the thickest was 32 mm at soil level in the same 
plant. Cortex was only 1.5 mm thick in the seedlings. The outermost cortex cells 
were columnar and aligned in palisades, the palisade cortex was about 3.5 mm 
thick. Cells of the inner cortex (located between the base of the palisade cortex 
and the phloem) consisted of large, rounded parenchyma cells. Cortical bundles 
were abundant throughout the inner cortex, extending to the base of the pali- 
sade cortex, and each bundle contained both xylem and phloem. Cortical 
bundles were easily visible by handlens and dissecting microscope. 

A slender pith was present in the center of all stems. It was only 1.5 mm in 


al 


diameter in seedlings, from 4 to 6 mm in plants of medium size, and 10 to 27 
mm in diameter in the largest plant (ie. plant *1214, which had the greatest 
girth). It consisted of just parenchyma cells with very rare spherical crystals 
and no mucilage. There were no medullary bundles at all and no dilated met- 
axylem. The lack of xylem in the pith was easily visible by handlens and dis- 
secting microscope: with both of these, shoot pith looked very clean and homo- 
geneous whereas the root's pith-like region was coarse and granular due to 
xylem in the dilatated region. 

Young shoots had a ring of collateral vascular bundles located between 
pith and cortex, older shoots had secondary xylem and phloem as well. Sec- 
ondary xylem in shoots was similar to root wood. Rays were narrow (149 sd 134 
um, just one or two cells wide) and consisted of parenchyma cells with no 
sclerification at all. The axial system consisted of small numbers of vessels and 
paratracheal parenchyma but large amounts of WBTs. As in roots, axial masses 
were narrow, only about 318 sd 179 um wide. No xylem fibers were present in 
any sample. Due to the narrow rays and axial masses and the lack of fibers, 
shoot wood resembled root and the two could not be distinguished if a micro- 
scope view contained only wood and no other tissues. Secondary phloem in shoots 
did not produce a cortex-like region as it did in roots; instead, as the sieve tube 
members stopped conducting, phloem collapsed into a thin, tangential band. 

Allsubterranean portions of Lophophora shoots were covered by bark simi- 
lar to that on older portions of roots. An unusual feature was that shoot bark 
occasionally contained crystals and vascular bundles, indicating that the cork 
cambium had arisen deeply enough in the shoot cortex to cut across cortical 
bundles; however, both crystals and vascular bundles were too small to be vis- 
ible with a handlens examination of bark. 


Structure of the hypocotyl 

The hypocotyl is the short (less than 10 mm long) region located between the 
seedling shoot (epicotyl) and the seedling root. The structure of the hypocotyl 
in L. williamsii had characters of both the root and shoot. The center of all hy- 
pocotyls was root-like because it consisted of dilatated metaxylem and inner- 
most secondary xylem, so it too was pith-like. It could be identified as not be- 
ing a true pith by the presence of vessel elements and WBTs interspersed with 
the parenchyma cells. The outermost regions were true cortex, and even though 
a hypocotyl is not a part of the shoot, the hypocotyls of Lophophora had corti- 
cal bundles. Hypocotyl cortex width was wider than that of the cortex-like 
region in roots, narrower than the true cortex of shoots in each plant. All hypo- 
cotyl samples had abundant secondary xylem and phloem, which was similar 
to that in both roots and shoots. Hypocotyl bark was similar to that of shoots, 
having occasional bits of cortical bundle that had been cut off by a cork cam- 
bium located deep within the cortex. 


586 BRIT.ORG/SIDA 22(1) 


Taste 2. Distinguishing characters of shoots and roots of L. williamsii. 


Shoots Roots 

Outer tissues True cortex; appears granular due to Cortex-like region; appears smooth 
presence of cortical bundles. At least due to lack of cortical bundles. At 
5mm or more thick. most only 3 mm thick. 

Center True pith; appears smooth due to Pith-like region; appears granular due 
lack of medullary bundles and lack of to dilatated metaxylem and innermost 
dilatated metaxylem. Width is not a secondary xylem. 
reliable criterion. 

Withered Sometimes present, not always easy Never present. 

tubercles to identify if bark is rough. 

Lateral roots Never present on shoots? May be Common on taproots, but could be 

confused with post-harvest absent. 


adventitious roots. 


DISCUSSION 

This study shows that roots and shoots of L. williamsii differ significantly in 
several features. At least in fresh plants, the two organs can be distinguished 
easily and reliably using just a handlens or dissecting microscope (Table 2). Both 
root and shoot have an outer region that resembles cortex, but the true cortex 
of shoots has a granular appearance because it contains numerous cortical 
bundles (Fig. 15), as is true of many cacti (Sajeva & Mauseth 1991; Mauseth & 
Sajeva 1992). In contrast, the outer region of roots resembles cortex but is in fact 
an accumulation of secondary phloem, which has a very smooth appearance 
as seen with a handlens. The vascular bundles of lateral roots pass through 
this cortex-like region of secondary phloem, but because lateral roots are so 
sparse and because their vascular bundles are oriented vertically, there is little 
chance of confusing the shoot and root outer tissues. Roots of other cacti also 
have this outermost cortex-like region (Stone-Palmquist & Mauseth 2002). The 
true cortex of the root of L. williamsii is pure parenchyma (Fig. 16) and does 
not contain secondary phloem. 

The center of shoots is occupied by true pith, which is homogeneous in 
appearance due to the lack of medullary bundles in Lophophora williamsii (Fig. 
17). Medullary bundles are common in many species of cacti but lacking in 
others (Mauseth 1993). In the roots of most species, metaxylem either has no 
parenchyma or if it does, the parenchyma does not undergo proliferation, so 
that roots of most plants have no pith-like region at all and can easily be distin- 
guished from shoots (Mauseth 1988). The pith-like region of roots in L. 
williamsit (Fig. 18) makes the roots look like shoots at first glance or with just 
the naked eye, but because the pith-like region in cacti originates by cell divi- 
sion in root metaxylem (Gibson 1978; Loza-Cornejo & Terrazas 1996), it has a 
granular appearance when examined with a handlens. 


— 


- 


L 
2 


oe rs ra Tk 4) 


Fic. 15. Cortex of shoot of L. williamsii. A the parenchyma. All visible 


parenchyma cells are cortex parenchyma. Scale bar (lower left) = 1 mm. 


Two characters might be useful for distinguishing shoots from roots with- 
out cutting plants open to examine the cortex-like regions and pith-like regions. 
Lateral roots emerged from the sides of other roots and from the sides of hypo- 
cotyls, but none was seen on any part of the three adult shoots we examined. It 
is possible that shoots had produced adventitious roots which had either bro- 
ken off when the plants were collected or which had abscised before collection. 
No remnants of such roots were seen when we examined the sides of subterra- 
nean portions of shoots witha dissecting microscope, but these plant parts were 
so wrinkled and bark-covered that we could have missed any that were present. 
However, as we examined the microscope slides of subterranean portions of 
shoots, we did not encounter any vascular bundles that would have indicated 
adventitious roots had been present. One plant, examined in the field, had a 
root emerging from its side at about the same level as a lateral branch; if that 
root was emerging from the shoot rather than the hypocotyl then shoots as well 
as roots might bear roots. Now that anatomical characters can be used to dis- 
tinguish roots from shoots, it will be possible to examine more plants in the 


588 BRIT.ORG/SIDA 22(1) 


All Wl + 1 } l Ah £ . in: i] 


Fic. 16. C f f L. williamsii 


Scale bar = 1mm. 


field to see how frequently shoots bear roots from their sides, and the extent to 
which such adventitious root development is associated with branching of sub- 
terranean stem tissue in response to peyote harvesting or removal of the apical 
meristem by natural processes. 

If the below-ground portion of the plant has withered tubercles, it must be 
part of the shoot rather than root or hypocotyl. However, we did not find any 
withered tubercles on one of our adult specimens despite a search with a dis- 
secting microscope. Apparently they either wither so much that they become 
unrecognizable or they abscise, so their absence cannot be used as proof that 
the structure is either hypocotyl or root. When trying to find withered tubercles 
in the field, one should search in a line continuous with the line formed by the 
rows of tubercles in the aerial shoot, because all tubercles are formed in rows 
Gust as on the ribs of columnar cacti: Mauseth 2000). 

If a plant remnant is to sprout and continue growing after its top has been 
harvested, the presence and health of these withered tubercles is important. If 
a plant is harvested by being cut too low, only root or hypocotyl will remain in 


=, 


Fic. 17. Pith of fst fl. williamsii 


} ‘ t4L n “aL + | a. 1inf+\ 
/ 


Arrows 
7 fal 1 L ij +4 rae 4 £L J} 4b Leal ‘aL 


. ul r ¢ é/ t Fr 
nol £tha ¢ lar hindloc tc ct feiahe) and all the call ho lof. sa = 
1 g I Scale bar = 1mm. 


the ground and neither of these have axillary buds, so neither can produce a 
bud to replace the harvested shoot. If the plant is harvested by being cut through 
the subterranean shoot, and if the remaining portion of shoot has healthy tu- 
bercles—withered but with an axillary bud—then the remaining portion should 
be able to sprout and grow and be ready for harvesting again in a few years. But 
if the remaining shoot has abscised all its tubercles, or if they have withered so 
much that they are no longer healthy, then the remaining piece of plant will 
not be able to sprout and will instead eventually die for lack of photosynthetic 
tissues. Tubercles located higher on the subterranean portion of the shoot are 
younger and presumably healthier than those lower down, deeper in the soil 
and closer to the root. If plants are harvested by cutting the subterranean shoot 
rather high—closer to soil level—the greater the chances are that the residual 
piece of plant will have healthy tubercles and will be able to re-sprout. 

With the information discovered in the current study, we now have the tools 
to examine in detail the potential for regrowth after different types of harvest- 


590 BRIT.ORG/SIDA 22 


1) 


3 7 t 7 


Fic. 18. Dilatated metaxylem of center of root of L. williamsii. All indicate masses of dark-staini | 
+ L 1 + All hL Ie? +h H 
? fe t 7 


not pith Scale bar= 1mm. 


ing. A set of plants could be harvested at various depths below soil level with 
the certainty that all had been cut high enough that some shoot tissue had been 
left on the remaining plant portion. The harvested top of each could be exam- 
ined for withered tubercles. Presumably if plants are cut so high that their har- 
vested tops have several recognizable withered tubercles near the cut, then the 
remaining portion also has at least a few tubercles and will sprout. But if plants 
are cut so low that the harvested tops have few or no identifiable withered tu- 
bercles near the cut—and if the cut does pass through shoot tissue and not root 
tissue—then probably the remaining shoot also has few or no tubercles capable 
of sprouting. It is even possible to try cutting the plants through the hypocotyl 
tosee if itis capable of forming adventitious shoot buds; that capacity is rare in 
hypocotyls but is known to occur in a few species. 

Preliminary data on plants under greenhouse conditions were collected on 
Il plants over a period of three years (not synchronically). Six plants were cut 
low (approximately 1.5 crown diameters below the base of the crown, and in 
every case at least 50 mm below the base of the crown). The distal subterra- 


nean portions of those plants were then observed for at least eight months. No 
new stem branches were observed on any of the six low-cut plants. The other 
five plants were cut high (at about the base of the crown, and in no case more 
than 5mm below the base of the crown) and similarly observed. One or more 
crown-bearing lateral branches from the decapitated subterranean stem were 
observed within five months on three of the five plants, and within eight months 
on all five high-cut plants. 

Future studies include a similar greenhouse experiment on regrowth, with 
substantially larger numbers of plants and with varying measured depths of 
cut expressed as a fraction (or multiple) of crown diameter, so that crown di- 
ameter could be used as a practical guide for harvesting peyote in the field. It 
will be noted whether the cut goes through root, stem, or hypocotyl. That titra- 
tion of the effect of depth of cut on regrowth in the greenhouse will be followed 
by a similar experiment conducted in the field, with individually identified 
and permanently marked plants. It is anticipated that results in the field may 
differ from greenhouse results, due to harsher environmental conditions in the 
field and the possibility that some of the smaller plants in the field may have 
been harvested previously, perhaps leaving less than the critical mass of sub- 
terranean stem tissue needed to regenerate viable photosynthetic stem tissue. 
These factors may affect regeneration and survivorship in all experimental 
groups. 


ACKNOWLEDGEMENTS 


We thank Phil Dering for the photo in Figure 2, and Shirley Powell for the pho- 
tos in Figures 13 and 14. Kendall Craig and Tim Parsons did minor miracles in 
adding needed layers to some photographs and retrieving high-resolution ver- 
sions from low-resolution derivative versions. Special thanks are due to Mike 
Powell for his ongoing support extended from the SRSC herbarium. 


REFERENCES 


ANDERSON, E.F. 1995. The “peyote gardens’ of South Texas: a conservation crisis? Cact. Succ. 
J.(Los Angeles) 67:67-73. 

Bravo, H. 1931. Nota acerca de la histologia del peyote, Lophophora willliamsii Lemaire. 
Anales Inst. Biol. Univ. Mexico 2:3-14. 

Gieson, A.C. 1978. Structure of Pterocactus tuberosus, a cactus geophyte. Cact. Succ. J. (Los 
Angeles) 50:41-43 

Janot, M.and M. Bernier. 1933. Essai de localisation des alkaloides dans le peyotl. Bull. Sci. 
Pharm. 40:145-153. 

Loza-Corneso and Terrazas. 1996. Bol. Soc. Bot. México 59:13-23. 

Mauseth, J.D., G. MonteNecro, and A.M. Watckowiak. 1985. Host infection and flower forma- 
tion by the parasite Tristerix aphyllus (Loranthaceae). Canad. J. Bot.63:567-581. 

Mausetn, J.D. 1988. Plant anatomy. Benjamin/Cummings. Menlo Park, California. 


592 BRIT.ORG/SIDA 22(1) 


Mausetn, J.D. and M. Saseva. 1992. Cortical bundles in the persistent, photosynthetic stems 
of cacti. Ann. Bot. 70:317-324. 

Mausetx, J.D. 1993.Medullary bundles and the evolution of cacti. Amer. J. Bot.80:928-932. 

Mauseth, J.D., ¥, Uozumi, BJ. PLemons, and J.V. LANbrum. 1995. Structural and systematic study 
of an unusual tracheid type in cacti.J.Pl.Res. 108:517-526. 

Mausetn, J.D. 2000. Theoretical aspects of surface-to-volume ratios and water-storage Ca- 
pacities of succulent shoots. Amer. J. Bot. 88:1107-1115. 

Mauseth, J.D. 2004.Wide-band tracheids are present in almost all species of Cactaceae. J. 
Pl.Res. 117:69-76. 

Moreno, S. 2005, For Native Americans, peyote is dwindling. Austin American-Statesman, 
Monday, September 19, 2005, p.B3. 

Rounier, A. 1927.Le peyotl—la plante qui fait les yeux emerveilles. Doin, Paris. 

Saleva, M.and J.D. MausetH. 1991. Leaflike structure in the photosynthetic, succulent stems 
of cacti. Ann. Bot. 68:405-411. 

ScHuttes, R.E. 1938. The appeal of peyote (Lophophora williamsii) as a medicine. Amer. 
Anthropol.,n.s. 40:698-715. 

STONE-Patmauist, M.and J.D. MauseTH. 2002. The structure of enlarged storage roots in cacti. 
Int. J. Pl. Res. 163:89-98. 

Texas DeparTMENT OF PusLic SArety. 2005. Peyote sales totals and distributors of Texas. Unpub- 
lished data. Austin, Texas. 


TAXONOMY AND CONSERVATION OF MEDICINAL PLANTS 
IN CANAL-IRRIGATED AREAS OF PUNJAB, PAKISTAN 


Khalid Faroog Akbar! Mohammad Athar' 


Department of Botan California Department of Food and Agricult 
Government Post-Graduate College 1220 N Street, Room 325 
Sahiwal, PAKISTAN Sacramento, California 95814, U.S.A. 
kezm@brain.net.pk atariq@cdfa.ca.gov 


ABSTRACT 
The taxonomic position of 131 medicinal plants from the canal-irrigated areas of Punjab belonging 
to 112 families and 52 genera is presented. ae life form, Bae used and pharmaceutical uses are 
also described. The most i ae family was 21 medicinal species, followed 


ae aceae, Asteraceae, Lamiaceae, Solanaceae (6 species each), Cucurbitaceae, Malvaceae, and 
Poaceae (5 species each). Life analysis of these medicinal plants indicates that all life forms are 
eee with the majority herbaceous. To conserve this natural wealth, an integrated strategy for 
both saving species and meeting the increased demand for maticeel ne is suggested. The main 


points of this strategy should include formulation of a comprehensive policy, increased ethnobo- 
tanical surveys and sustainable production of medicinal ae through their cultivation. 


Beer ] ] | Ms H | 
[ r 


Key Worps: Canal irrigation, conservation, Pakistan 


RESUMEN 


Se presenta la posicion taxonémica de 131 plantas medicinales de las areas irrigadas del canal del 
Punjab aie a 52 familias y 112 géneros. Se describen también las formas vitales, partes 


laa) 
~ 
Q 
oO 
Lo) 
= 
Q 
(e) 
yn 
(** 
pS) 
S 
= 
5 
~ 
wn 
ge) 
io} 
fa 
pst) 
3 
> 
oO 
or 
wn 
rT 
~ 
ion 
p 
Q 
Oo 
ist) 
far) 
Q 
oO 
i=} 
ine) 
te 
oOo 
wn 
a] 
oO 
(@ 
oO 
n 
=| 
o 
a: | 
a 
3 
lean 
oOo 
wn 


usadas y u 
seguida por se Asteraceae, Lamiaceae, Solanaceae (6 especies cada una), Cucurbitaceae, 
Malvaceae, y Poaceae (5 especies cada una). El analisis de las formas vitales de estas plantas 
medicinales indica que estan representadas todas con predominio de las herbaceas. Para conservar 
esta riqueza Batra se ak ere una esttaicely ion tanto poe janes ie Src como Bal a 


abordar | | I I 
] ] p> ies a }; 1 auc 
ir ncluir la for mulaci la produccion 


sostenible de plantas medicinales eden su cultivo. 


INTRODUCTION 


Pakistan has the distinction of having the largest contiguous gravity-flow irri- 
gation system in the world. The Pakistani irrigation system has been greatly 
expanded since independence in 1947. This irrigation system serves as a life- 
line for sustaining agriculture and is the main source of livelihood for the 70% 
population. Punjab has a well-developed irrigation system with a net work of 


In; 


Disclaimer: The views expressed in this articles are ee of authors and do not necessarily aes nt those of 


their Bene departments. The publication o endorse- 
ment of the use of these plants as herbal medicine. The authors or their departments make no warranty, ex- 


ee or implied, and assume no legal liability for the use of these plants for medical purposes 


SIDA 22(1): 593 — 606. 2006 


594 BRIT.ORG/SIDA 22(1) 


— 


canals, water channels and distributaries. The canal-irrigated areas account for 
90% of agricultural production and employ 54% of the labor force (Khan 2002). 

Pakistan is rich in plant resources with more than 6,000 plant species o 
higher plants (Nasir & Rafiq 1995). The irrigated areas of Punjab are rich in 
phyto-diversity, containing a wealth of cultivated crops, natural vegetation and 
wild flora (Ahmad 2003). This includes plants used in traditional herbal medi- 
cine. Research on medicinal plants in the irrigated areas of Punjab has been 
generally ignored in respect to developing a scientific understanding of the 
dynamics of natural resources, organization and evolution of human systems. 
This apparent lack of attention to irrigated, cropped areas has resulted ina pau- 
city of scientific data about the natural vegetation of these areas. There are a 
few studies listing the medicinal plants from various regions of Pakistan (Ah- 
mad & Waseem 2004; Athar & Ahamd 2004; Athar & Siddiqi 2004; Gill 2003). 
However, the irrigated areas of Punjab have not been explored adequately in 
floristic surveys, and a comprehensive documentation of the medicinal flora 
of the region also is lacking. Pressure on the wild plants of these areas is ex- 
pected to increase with increasing agricultural, urban and industrial activities. 
The natural vegetation, including medicinal plants, of these areas is severely 
degrading, and there is an immediate need to protect and preserve these plant 
resources. This study is aimed at compiling a list of medicinal plants in canal- 
irrigated areas of Punjab with information about their taxonomic position, life 
form and pharmaceutical utilization of different plant parts. The study also 
explores the reasons for the decline of medicinal plants in the irrigated areas 
and suggests measures for their conservation. 


rH 


je 


MATERIALS AND METHODS 


A literature search was conducted to determine the medicinal plants used for 
various disorders or diseases in canal-irrigated areas of Punjab, Pakistan (Baquar 
1989, 1995; Joshi, 2003; Maheshwari 2003; Nasir & Ali 1972; Nasir & Rafiq 1995; 
Palaniswamy 2003; Rahim 1996; Rizvi 1998: Shaheen et al. 2003; Tivari & 
Tandon 2004; William & Ahmad 1999). Their taxonomic position, life form and 
pharmaceutical utilization of different plant parts were also determined. The 
plants included herbs, shrubs and trees. The genera were arranged alphabeti- 
cally within each family. The English and local names that are commonly as- 
sociated with these plants are provided. The nomenclature and classification 
followed Nasir and Ali (1972), and author citations followed Brummitt and Pow- 
ell (1992). 


RESULTS AND DISCUSSION 
This study represents the first comprehensive survey of medicinal plants in 
the irrigated areas of Punjab, Pakistan with suggested strategies to conserve this 


natural wealth by both saving the medicinal plants and meeting their increased 


TaBLE 1.Some important medicinal plants in irrigated areas of Punjab. 


Species Local name English name Life form Part used Pharmaceutical uses 
MONOCOTYLEDONS 

Agavaceae 

Agave americana L. Banskeora Agave Herb Leaves, roots Astringent 

Alliaceae 

Allium cepa L. Onion Herb Bulbs, leaves, seeds Anti-ulcera 

Allium sativum L Lehson Garlic Herb Bulbs, leaves eae a pressure 

Asparagaceae 

Asparagus adscendens Roxb. Musli sufaid Garden asparagus = Shrub Seeds, roots Antidiarrhoeal, demulcent 
galactogogue 

Asphodelaceae 

Aloe vera (L.) Burm. f. Gheekawar Aloe Herb Whole plant Cosmetic agent, febrifuge, 
anthelmintic 

Cyperaceae 

Cyperus rotundus L. Dela Sweet cyperus Herb Roots Diaphoretic, lithontriptic, liver tonic 

Poaceae 

age lias (L ee Tabasheer Bamboo Tree Leaves, roots Tonic, astringent antidiarrhoeal 

Ousa 5 (L.)Vo Bamboo Tree Leaves, stem Styptic, tonic, desiccative 
ymbopogon citratu Izkhir Russ grass Herb Stem Antiseptic, stomachic, styptic 

(DC.) Stapf. 

Cymbopogon jwarancusa Khowi ghas Russ grass Herb Flowers Detoxifier, astringent and tonic 

(Jones) Schult. 

Zea mays L. Makai Maize, corn Herb Flowers Astringent, chloretic, diuretic and 
remedy for urinary infection 

DICOTYLEDONS 
Amaranthaceae 
Achyranthes aspera L. Pathkanda Prickly chaff Herb Whole plant Astringent, diuretic 


NVLSINVd ‘ 


IVNIDIGUW GVAly ONY avadny 


965 


Tasce 1. continued 


Species Local name English name Life form Part used Pharmaceutical uses 

Apiaceae 

Apium g lens L. Soya Dill Herb Roots, seeds, Rheumatism, arthritis, indigestion 

stem flatulence, urinary t 

inflammation and on 
neurasthenia, sleeplessness, anxiety 
and nervous breakdown 

Centella asiatica (L.) Urb. Brahmi booti Pennywort Herb es Anti-inflammatory, diuretic 

Coriandrum sativum L. Dhania riander Herb Leaves, stem Antispasmodic, appetizer, aromatic 

Daucus carota L Gajar Carrot Herb Roots, stem Aphrodiasic, exhilarant, diuretic 

Foeniculum vu inate Mill. Saunf Fennel Herb Leaves, stem Digestive, lactagogue, masticatory, 
flavoring agent 

Trachyspermum ammi (L.) Ajowan Omum Herb Leaves, stem Stimulant, antispasmodic 

Sprague ex Turrill 

Apocynaceae 

Alstonia scholaris (L.) R. Br. Shaitan-ka- Devil's tree Tree Bark Fever treatment 

Jhad 

Nerium oleander L. Kaner Oleander Shrub Leaves, roots Abortifacient, attenuant 
anti-inflammatory 

Tabernaemontana divaricata Chandani Nero's crown Herb Flowers, roots Anthelmintic 

(L.) R.Br.ex Roem. & Schult. 

Asclepiadaceae 

Calotropis gigantea (L.) Madar Gigantic Shrub Whole plant Expectorant, alterative 

W.T. Aiton swallowort 

Calotropis procera (Aiton) Ak Swallowort Shrub Whole plant Diaphoretic, emetic, diuretic 


W.T. Aiton 


96S 


(L)@2 VaIS/D¥O' LIYE 


Taste 1.continued = 
=> 
= 

Species Local name English name Life form Part used Pharmaceutical uses S 
= 

Asteraceae ai 

Ageratu nyzoides L. Ajganda Herb Flowers, leaves Emetic, digestive tonic = 

lendula officinalis L. Zergul Calendula Herb Flowers Diaphoretic, antiemetic = 

Carthamus tinctorius L. Kusumba Safflower Herb Flowers, stem mmenagogue, ae sedative, = 

stimulant 

Eclipta prostrata (L.) L. Bhangra Eclipta Herb Whole plant Deobstruent, antiasthmatic 

atricaria recutita Babuna Chamimila Herb Carminative, stimulant 

Xanthium strumarium L. Banukra Cocklebur Herb Whole plant Diaphoretic, emollient 

Boraginaceae k 

Cordia dichotoma G. Forst. Lasora Sebestan plum Tree Fruits Demulcent, laxative = 

Heliotropi Oont chara Herb Whole plant Skin disorders = 

Brassicaceae 

Brassica juncea (L.) Czern. Rai Brown mustard Herb Seeds Emetic 

Lepidiu tivu Chandrasur Common cress Herb Leaves, roots Stimulant, anti-inflammatory 

Sisymbrium irio L. Khub kalan Hedge mustard Herb Leaves, seeds Skin disorders 

actaceae 

Opuntia dillenii (Ker Gawl,) Chappal Prickly pear Shrub Whole plant Demulcent, expectorant 

Cannaceae 

Canna ind Aqeeq Indian Shot Herb Roots, seeds Diaphoretic, diuretic, demulcent 

annabinaceae 

Cannabis sativa L. Bhang Indian hemp Herb Flowers, leaves Sedative, narcotic 

Capparidaceae 

Capparis spinosa L. Karir Capper plant Shrub Roots Anti-inflammatory, anti-flatulent, 

resolvent 
Cleorne gynandra L. Huinul Caravalia seed Shrub Leaves, seeds Rubefacient, anthelmintic 
Cleome viscosa L. Hurhur Wild mustard Herb Leaves, seeds Rubefacient, anthelmintic 


£6S 


Tage 1.continued 


Species Local name English name Life form Part used Pharmaceutical uses 

Chenopodiaceae 

Chenopodium alb Bathu White goose foot Herb Seeds Anthelmintic 

phania amb d Sak Mexican tea Herb Seeds Vermifuge 

Mosyakin & Clements 

Combretaceae 

Terminalia arjuna (Roxb. ex Arjan Arjun tree Tree Bark Stomachic, anti-ischemic and 

oe ) vee & a dla abcae iv 

a (Gaertn.) Bahira Beleric myrobalan Tree Fruits Bron suilstany antispasmodic 

poe ot gmatic, expectorant and 
sedative activities 

Terminalia chebula Retz. Harir Bidda nut Tree Fruits Stomachic, alterative 

Convolvulaceae 

Cuscuta reflexa Roxb. Akas bel Dodder Herb Seeds Carminative, alterative, anodyne 

Crassulaceae 

siete oe (Lam.) Pers. Zakhame hayat Life plant Herb Leaves Antiseptic, styptic 

urbitacea 

as co ee (L.) Tumma Colocynth Herb Fruit, roots Anthelmintic, antipyretic 

Schrad. 

Citrullus lanatus (Thunb.) Tarbooz Water melon Herb Fruits, seeds Laxative, brain tonic 

Matsum. & Nakai var. /anatus 

Lagenaria siceraria (Molina) Kaddo Gourd Herb Seeds Brain tonic 

andl. 
ita vetnangula (L ’ ‘a Ghia tori Herb Fruits Bitter tonic, diuretic 
Karela Bitter gourd Herb Fruits, a seeds Anthelmintic, purgative 

Euphorbiaceae 

Chamaesyce hirta (L.) Millsp. Dhodak Snake weed Herb Leaves, stem Purgative, rubefacient 

Euphorbia neriifolia L. Thanda thor Milk bush Tree Latex, roots 


Purgative, expectorant, 
antispasmodi 


865 


(L)@Z VaIsS/OYO'LINA 


TaBLe 1. continued 


Species Local name English name Life form Part used Pharmaceutical uses 
Phyllanthus niruri L. Bhoin avail Myroblan Herb Leaves, roots Diuretic, astringent, bitter tonic 
Ricinus communis L. Arind Castor oil Shrub Leaves, seeds Emetic, anti-inflammatory 
Fabaceae (Caesalpinioideae) 
Bauhinia purpurea L. Kaliar Purple Bauhinia Tree Bark, flowers, Purgative, indigestion, body pain 
flower buds 
Bauhinia semla Wunderlin Kandla Bauhinia Tree Bark, leaves, Astringent, diarrhea, dysentry 
flowers 
Bauhinia variegata L. Kachnar Mountain ebony Tree Bark, flower Blood purifier, anthelmintic 
5, roots 
Cassia fistula L. Amaltas Golden shower Tree Fruits, leaves, roots Emollient, febrifuge 
ista absus (L.) H.S. Chaksu Chaksu Herb Leaves, seeds Astringent, detersive, heamotonic 

Irwin & Barneby opthalmic 
Senna alata (L.) Roxb. Dadmurdan Ringworm shrub Tree Leaves Purgative, expectorant, aperient 
Senna alexandrina Mill n Senna Herb Leaves Purgati 
Tamarindus indica L. Imli Tamarind Tree Bark, leaves, fruits ooling, antibilious 
Fabaceaae (Mimosoideae) 

cacia modesta Wall. Phulai Tree Gum Emollient 
Acacia nilotica (L.) Delile Kikar Acacia Tree Flowers, leaves Emollient, astringent, styptic 

Albizia iter .) Benth. Siris Lebbek tree Tree Bark, leaves Astringent, oral therapy 
Mimosa p Chhui-mooi Touch me not Herb Leaves Emetic, renal diseases 
pene pee 

Alhagi maurorum Medik. Jawansa Camel's thorn Shrub Whole plant Laxative, demulcent, expectorant 
Clitoria ternatea L. Gokarni Butterfly pea Herb Roots, seeds Cathartic, dem 
Cullen plicata (Delile) Bakuchi Babchi Herb Seeds Anthelmintic, ee 
C.H. Stirt 
Erythrina stricta Roxb. var. Pangra Coral tree Tree Bark Cathartic, anthelmintic 


suberosa (Roxb.) Niyomdham 


WNDIGIW aVAly ONY dVaAy 


NVISDIVd 


665 


Tasle 1. continued 


Species Local name English name Life form Part used Pharmaceutical uses 
Indigofera tinctoria L Nee| Indigo plant Herb Whole plant Deobstruent, alterative 
Sesbania grandiflora (L.) Bansa Sesbania Tree Roots, flowers Rheumatism, fever, diabetes 
Per 
eee sesban (L.) Merr. Janter Sesbania Shrub Leaves, flowers Suppuration, anti-fertility 
Trigonella foenum-graecum L Methi Fenugreek Herb Leaves, seeds Aperient, nutritive, expectorant 
Vigna unguiculata (L.) Walp. Kulatha Horse gram Herb Seeds Astringent, aiere lithontriptic 
subsp. cylindrica (L.) Verdc. 
Lamiaceae 
Lallemantia royleana (Benth. Tukham-i- Herb Seeds Cooling, mucilaginous 
Benth. balanga anti-inflammatory 
Mentha arvensis L. din Mint Herb Whole plant Carminative 
Mentha xpiperita L. Podina Peppermint Herb Leaves Aromatic oil, digestive aa 
Ocimum basilicum L Niazb Sweet basil Herb Flowers, seeds Carminative, expectoran 
Ocimum tenuiflorum L. Tulsi Holy basil Herb Flowers, leaves, Stimulant, antiseptic 
roots 

Salvia plebeia R. Br. Salbia Sage Herb Seeds Treating boils and wounds 
Linaceae 
Linum usitatissimum L Alsi Linseed Herb Seeds Diuretic, emollient 
Lythraceae 
Lawsonia inermis L. Mehndi Henna plant Shrub Whole plant Hair tonic, hepatic disorders 

lvaceae 
Abutilon bidentatum A. Rich. Pattaka Country mallow Herb Leaves, seeds Demulcent, diuretic 
Abutilon indicum (L.) Sweet Peeli booti Country mallow Shrub Leaves, roots, seeds Demulcent, laxative, sedative 
Gossypium herbaceum L Kapas Cotton erb eeds Detersive, emollient, expectorant 
Malva sylvestris L. ee Common mallow Herb Fruits, leaves Throat infections 
Sida cordifolia L. mak Country aa Shrub Leaves, roots Astringent, diaphoretic 


009 


(L)@Z ¥QIS/9YO LYS 


Table 1.continued 


Species Local name English name Life form Part used Pharmaceutical uses 

Meliaceae 

Azadirachta indica A. Juss. Neem Margosa tree Tree Whole plant Adrenalgic stimulant, dyspepsia, 
skin ae insecticidal 

Melia azedarach L. Drek Persian lilac Tree Flowers, leaves Anthelm antispasmodic 

Toona ciliata M.Roem. Tun Indian mahogany Tree Bark ern 

Moraceae 

Ficus benghalensis L. Bar Banyan Tree Leaves, latex Astringent, viscous, resolvent 
desiccative 

Ficus religiosa L. Pipal Pipal tree Tree Bark, fruits Astringent, antiasthmatic 

Ficus virens Aiton Pilkhan Tree Fruits, stem Astringent, stomachic 

ringaceae 

Moringa oleifera Lam. Soanjan Horse radish tree Tree Flowers, leaves Anthelmintic, antipyretic 

Myrtaceae 

Barringtonia acutangula (L. Jugar Indian Oak Tree Bark, roots, seeds Astringent, emetic 

Gaertn 

Mine communis L. Wilayti Myrtle Shrub Fruits, leaves Antiseptic, disinfectant 

mehndi 

Psidium guajava L. Amrood Guava Tree Fruits, seeds Astringent, febrifuge, antiseptic, 

laxative 
taginaceae 

Se diffusa L. Itsit Hog weed Herb Leaves, roots Diuretic, diaphoretic, anthelmintic 

Mirabilis jalapa L. Gul-i-abbas Four o'clock plant Shrub Leaves, roots Demulcent, antispasmodic 

Nymphaeaceae 

Nelumbo nucifera Gaertn. Kanwal Water lily Herb Fruits, leaves Cardiac tonic, demulcent 

Nymphaea lotus L. Nilofar White lotus Herb Flowers Cardiac tonic, hepatic disorders 


IWNDIGIW UVALY ONY ava 


NVLSIAVd 


L09 


TasB_e 1. continued 


709 


Species Local name English name Life form Part used Pharmaceutical uses 

Oleaceae 

lasmi inale L. Chambeli Jasmine Shrub Flowers, leaves Anthelmintic, diuretic 
Nyctanthes arbor-tristis L. Kuri Night jasmine Shrub Leaves, seeds Antibilious, laxative 

Oxalidaceae 

Oxalis corniculata L. Surchi Indian sorrel Herb Leaves Anti-scorbutic, refrigerant 
Papaveraceae 

Argemone mexicana L. Stayanasa Yellow thistle Herb Whole plant Alterative, stimulant 

Pedaliaceae 

Sesamum indi Til Sesame Herb Seeds Aphrodiasic, fattening, nourishing 


Plantaginaceae 
Plantago ovata Forssk. lsabaghol Plantain Herb Bark, seeds Cooling, mucilaginous 
anti-inflammatory 


Plumbaginace 
Plumbago ee L. Chitrak Lead wort Herb Roots Stimulant, antiseptic, detersive 
refrigerant 


Portulacaceae 


Portulaca oleracea L. Dholica Purslane Herb Seeds Demulcent, diuretic, astringent 
Ranunuculaceae 

Nigella sativa L. Kalonji Black cumin Herb Seeds Diuretic, galactagogue, stimulant 
Rl amnaceae 

phus jujuba Mill. Beri Jujube Tree Leaves, seeds Anti-inflammatory 
Rutaceae 
Aegle marmelos (L.) Corréa Bel Bil Tree Flowers, leaves Laxative, expectorant 
Salvadoraceae 

lvad leiodes Decne. Pil Tooth brush tree Tree Bark, leaves, seeds Vesicant, stimulant 
Salvadora persica L. Pilu Mustard tree Tree Fruits, leaves Diuretic, anthelmintic 


(L)@Z ¥OIS/9YO'LIYS 


Taste 1. continued 


Species Local name English name Part used Pharmaceutical uses 
Sapindaceae 
Cardiospermum hali. Kanphuti Baloon vine Whole plant Demulcent, laxative 
Scrophulariaceae 
Bacopa monnieri (L.) Pennell. Brahmi Thyme Gratiola Whole plant Febrifuge, nervine 
Simaroubaceae 

Ailanthus excelsa Roxb. Maharuk Talbot Bark, leaves Astringent, febrifuge 
baaanee 

Mirch Red pepper Leaves, seeds Pungent, sialgogque 
Datura metel L. Datura Apple of Peru Leaves, seeds Narcotic, toxic, antispasmodic 
Tamatar Tomato Fruits Antioxidant, carminative 

Sol anum dete L. Mako Black nightshade Roots, seeds Cardiac tonic, diaphoretic 
Solanum virginianun Kandiari Indian Solanum Whole plant Expectorant, diuretic 

Withania somni ae Asgandh Winter cherry Whole plant nthelmintic, stimulant 
Tiliaceae 
Corchorus olitorius L. Kost Jute plant Seeds, leaves Carminative, astringent 
Grewia asiatica L Phalsa Grewia Fruits Carminative 
Zingiberaceae 
Curcuma longa L. Haldi Turmeric Seeds Antimutagenic, antioxidant 

y hyllaceae 
Fagonia cretica L. Prickly cover eaves Refrigerant, febrifuge 
Peganum harmala Harmal Harmal Roots, seeds Narcotic, hypnotic, emetic 
Tribulus terrestris L. Pakhra Puncture vine Aphrodisiac, diuretic 


TWNIDIGIW UVALY ONY avaaAy 


NVLSTAWd 


£09 


604 BRIT.ORG/SIDA 22(1 


demands in the traditional health care system. A total of 131 medicinal plant 
species in 112 genera belonging to 52 plant families are reported from the ca- 
nal-irrigated areas of Punjab (Table 1). It is interesting to note that most of the 
medicinal plants were Dicots (48 families), while Monocots constitute only a mi- 
nor fraction of them (3 families). The most important family was Fabaceae con- 
taining 21 medicinal species. This was followed by Apiaceae, Asteraceae, 
Lamiaceae, Sol (6 species each), Cucurbitaceae, Malvaceae, and Poaceae 
(5 species each). Life form analysis of these medicinal plants indicated that all 
life forms were represented, but the majority belonged to herbaceous forms 
(Table 1). Herbs (80 species) were found to be the most common life form fol- 
lowed by trees (32 species) and shrubs (19 species). Most frequently utilized 
plants parts included leaves (54 species), seeds (44 species) and roots (27 spe- 
cies). Whole plants of 18 species were utilized in herbal medicine. The bark, 
fruits and flowers (16 species each) and stem and branches (10 species) also 
served as medicines. Onion and garlic bulbs and gums and latexes from some 
of the plants were also used as medicines. 

The biological and physical changes associated with fragmentation and 
loss of habitats and canal irrigation have many residual and secondary effects 
that aggravate the problem of phyto-diversity loss. These include pollution, 
decline of soil fertility, water logging, the salinization or acidification of soil 
and water, the siltation of water channels and increased desertification. Among 
these, salinity and water logging of soils in Punjab are the most important prob- 
lems (Davidson 2000). Saline soils, high temperatures and increased rate of 
evaporation, and limited water supply pose a serious limit back on the growth 
and distribution of native plants (Hussain et al. 2003). Desertification is an- 
other threat to native plants of Punjab because it particularly affects the biota 
of areas with 200-800 mm of average rainfall (Khan 2000). The activities that 
lead to or enhance desertification include cultivation of unsuitable terrains and 
soils, over-grazing and lack of proper livestock management practices, and ex- 
cessive removal of shrubs and trees for fuel wood (Ahmad & Waseem 2004). 
This has resulted in the loss of many herbaceous species, including medicinal 
plants, in many areas of Punjab (Khan 2000). 

The irrigated areas of Punjab are as rich in phyto-diversity and deserve all 
possible protection and conservation measures to prevent loss of wild plants. 
The conservation and sustainable management of medicinal plants requires a 
strategic action at different levels. As part of the conservation of medicinal 
plants, the management and use of agricultural lands need major revamping. 
Farmers should realize that their methods of land use and farming practices 
affect the surrounding environment and the organisms in that environment. 
Asa first step towards better land management, clearing and fragmentation of 
native vegetation should be stopped and the extent of native plant reserves 
should be increased. 


os 


may 


AKBAR AND ATHAR, MEDICINAL PAKISTAN 605 


Medicinal plants are one of the few natural products of developing coun- 
tries that could be marketed at higher prices (Athar @ Ahmad 2004; Shaheen 
et al. 2003). The ever growing market and demand for herbal ingredients cre- 
ates possibilities for cultivation of medicinal plants as crops (Hussain et al. 
2003). Such efforts could provide a substantial supply of herbal plants and help 
increase the income of the farmers. With continuous increase in use and de- 
mand for herbal products, the gap between demand and supply is widening 
(Crockett & Khan 2003). It is estimated that world trade in medicinal plants 
and related products may reach to US$ 5 trillion by 2050 (Purohit & Vyas 2005). 
Most of the demand so far has been met from wild sources, which cannot last 
much longer. Since medicinal plants are a source of additional earnings and 
traditional treatments for a major part of rural population, it is recommended 
that these plants may be propagated and domesticated as cash crops. Thus, cul- 
tivation of medicinal plants could be a potential source of economic develop- 
ment that would also meet conservation objectives. 


ACKNOWLEDGMENTS 


Special gratitude is expressed to Joseph H. Kirkbride, USDA, Agricultural Re- 
search Service, Beltsville, MD, for his help in checking the nomenclature and 
taxonomy of the plants and M. Akmal Siddiqi, Marshfield Medical Research 
Foundation, Marshfield, WI, USA for valuable suggestions and helpful criticism 
on the manuscript. The authors are also grateful to M. Iqbal Choudhary, H.EJ. 
Research Institute of Chemistry, University of Karachi, Karachi, Pakistan for 
providing many original reprints used in the preparation of this paper. 


REFERENCES 


AHMAD, H. 2003. Capacity building for cultivation and sustainable harvesting of medicinal 
and aromatic plants. In: H. Anmad and A.A. Khan, eds. Conservation and sustainable 
uses of medicinal and aromatic plants of Pakistan. WWF-Pakistan. pp.32-37. 

Auman, H. and M. Waseem. 2004. Conservation status of some medicinal plants of the salt 
range. Zonas Aridas 8:1-8. 

AtHar, M.and Z. AHmab. 2004. Taxonomy, distribution and medicinal uses of medicinal le- 
gume trees of Pakistan. Sida 21:951-962. 

AtHar, M.and M.A. Sippial. 2004. Some reflections on the taxonomy of medicinal flowers of 
Pakistan. Sida 21:357-369. 

Baauar, S.R. 1989. Medicinal plants and poisonous plants of Pakistan. Printas, Karachi, Pakistan. 

Baauar, S.R. 1995. Trees of Pakistan. Royal Book Company, Karachi, Pakistan. 

Brummit, K.R. and C.E. Powe t (eds.). 1992. Authors of plant names. Royal Botanic Gardens, 
Kew, UK. 

Crockett, S.L. and |.A. KHan. 2003. Challenges of standardization: Marker compounds in 
plants species related and unrelated to top-selling herbs. J. Herbs, Spices & Med. Plants 
10:13-24. 


606 BRIT.ORG/SIDA 22(1) 


Davioson, A.P, 2000. Soil salinity, a major constraint to irrigated agriculture in the Punjab 
region of Pakistan:Contributing factors and strategies for amelioration. Amer.J.Alternat. 
Agric.15:154-159 

Git, M.A. 2003. Cultivation of medicinal and aromatic herbs: experiences of IMHC. In: H. 
Ahmad and A.A. Khan, eds. Conservation and sustainable uses of medicinal and aro- 
matic plants of Pakistan. WWF-Pakistan. Pp.23-31. 

Hussain, N.,A.R. Naseem, G. Sarwar, F. Muuees, and M.Jamit 2003.Domestication and cultivation 
of medicinal crops on salt-affected soil. In: H. Ahmad and A.A. Khan, eds. Conservation 
and sustainable uses of medicinal and aromatic plants of Pakistan. WWF-Pakistan. Pp 
38-44. 

JosHi, S.J. 2003. Medicinal plants. Oxford & IBH Publishing Company, New Delhi, India. 

KHAN, A.U. 2000. History of decline and present status of natural tropical thorn forest in 
Punjab, Pakistan. Biol. Conserv. 63:250-260. 

KHAN, F.K. 2002. A geography of Pakistan. Oxford University Press, Karachi, Pakistan. 

ManesHwarl, J.K. 2003, Ethnobotany and medicinal plants of Indian Sub-continent. Scien- 
tific Publishers, Jodhpur, India 

Nasir, E.and S.|. Au. 1972.An annotated flora of West Pakistan. Stewart Herbarium, Gordon 
College, Rawalpindi, and Department of Botany, University of Karachi, Pakistan. 

Nasir, YJ. and R.A. Raria. 1995. Wild flowers of Pakistan. Oxford University Press, Karachi, 
Pakistan. 

PaLaniswamy, U.R.2003.A guide to medicinal plants of Asian origin and culture.CPL Scien- 
tific Publishing Services Limited, Newbury, UK. 

PUROHIT, S.S.and S.P. Vyas 2005.Medicinal plant cultivation: A scientific approach. Agrobios, 
India. 

Rani, A. 1996. Our useful plants. Polymer publishers, Lahore, Pakistan, 

Rizvi, M.A. 1998. Medicinal flowers of Pakistan, Part-Ill. Horticultural Society of Pakistan, 
Karachi, Pakistan. Pp.48-52. 

SHAHEEN, F., M. ATTAUR RAHMAN, K.VASISHT, and M. lagat CHoubHaRY. 2003. The status of medicinal 

and aromatic plants in Pakistan, In: Medicinal plants and their utilization, UNIDO. Pp. 

7 Vy 


os. 


Tivari, M.and V. Tanbon. 2004. Medicinal plants. sha Books, New Delhi, India. 
Wiuam, J.T.and Z. AHmab. 1999, Priorities for medicinal plants research and development 
in Pakistan. Published by MAPPA, India, and IDRC, Canada. 


NOTES ON THE LIFE AND WORK OF 
JAMES BRIGHAM MCFARLIN, FLORIDA BOTANIST 


Tom Palmer 
1805 26" St.NW 
Winter Haven, Florida 33881, U.S.A. 
ABSTRACT 
James Brigham McFarlin contributed to Florida botany and his work provided a rare glimpse into 
the diversity that once existed in the central part of the state three-quarters of a century ago. 
RESUMEN 


ae ee Mean aaah a la botanica de Florida y su obra dio una ojeada rara a la 
Florida central hace 75 anos. 


James Brigham McFarlin (1901-1969) was among a small group of botanists 
whose work contributed to a better understanding of Florida’s flora through 
explorations and collections during the first half of the 20th Century. Unlike 
J.K.Small and other prominent botanists who visited Florida to collect and ex- 
plore, Jim McFarlin (Fig. 1) was a Florida resident. He used his familiarity with 
the area—particularly the Lake Wales Ridge in the central peninsula—his en- 
thusiasm and a keen eye to collect and record specimens that he regarded as 
new species or new forms of known species as well as species that were previ- 
ously undocumented in the state. Apart from a brief mention in Wunderlin et 
al. (2003), nothing appears to have been written about McFarlin’s life and work. 
This paper is an attempt to fill that gap. 

Jim McFarlin was born November 24, 1901, in New York City. He attended 
Mercersburg Preparatory School in Mercersburg, Pennsylvania and went on to 
attend Syracuse University, where he received a degree from the New York State 
College of Forestry in 1925. He then joined his parents, Robert Roy and Grace B. 
McFarlin, in Winter Haven, Florida. The elder McFarlins had moved to Winter 
Haven in 1918 (Burr 1974), where the elder McFarlin had purchased land and 
operated a citrus grove. After moving to Winter Haven, Jim McFarlin was presi- 
dent and manager of Winter Haven Ornamental Nurseries east of Winter Ha- 
ven (City Directory 1928). However, he was not a particularly successful busi- 
nessman and survived at times only through his father’s financial assistance 
(M. Bryant, pers. comm.). 

His primary interest at that time was botany, not the nursery business. He 
followed that interest from 1930 to 1933 with an assistantship from the Univer- 
sity of Michigan, where he pursued a doctorate. His graduate assistant stipend 
was $600 a year (Cherie Peterson, pers. comm.). His thesis work involved exten- 


SIDA 22(1): 607 - 613. 2006 


608 BRIT.ORG/SIDA 22(1) 


Fic. 1. James Brigham McFarlin (front left). 


sive collecting in Polk County and some surrounding areas in connection with 
the preparation of his dissertation (McFarlin 1935). No degree was awarded and 
his assistantship was not renewed after 1933. 

By 1933 McFarlin was married with one child, unemployed, and in the 


PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 609 


depths of the Depression. He found temporary work with the U.S. Department 
of the Interior's Civilian Conservation Corps. He was employed initially in the 
Chippewa National Forest, Cass Lake, Minnesota. In 1934 he was transferred to 
the CCC camp near Sebring, Florida, where work was underway to develop 
Highlands Hammock State Park, one of Florida’s first state parks. McFarlin be- 
came the botanist at the Highlands Hammock Botanical Garden and Arbore- 
tum and served on the Executive Committee of its Botanical Garden Associa- 
tion (Altvater 1966). He was recommended for that position by Harley Harris 
Bartlett, then chairman of the Department of Botany and director of the Bo- 
tanical Garden at the University of Michigan (letter, 17 February 1934, to Harry 
Lee Baker, Florida state forester). Dr. Bartlett had been a member of McFarlin’s 
graduate committee at Michigan. 

McFarlin was responsible for preparing some of the early interpretative 
materials for Highlands Hammock State Park (McFarlin 1939). Some of his field 
notebooks containing notes on specimens he collected during his field work in 
Florida were found around 2000 during an inventory of historical artifacts at 
the park, but their importance was not recognized until 2005. The pages in the 
notebooks are being digitally copied for use in a planned exhibit at the park’s 
CCC museum (D. Smith, pers. comm.). 

McFarlin lived in the Sebring area until at least 1942, but later moved back 
to Winter Haven (Burr 1974) and lived there until moving to Bradenton in 1949, 
shortly after the death of his father in October 1948. In Bradenton, probably 
because of the inheritance he received from his father’s estate, he was able to 
resume his career in the nursery business. He operated Begonia Gardens there 
until his death on March 10, 1969 (Anonymous 1969). According to his obituary, 
he and his wife Marie, who he married in 1927, had a son, James R. McFarlin, a 
daughter, Sarah Wells, and four grandchildren. 

McFarlin began his botanical collection as early as 1923, according to cor- 
respondence between 1924 and 1935 with J.K. Small (Florida State Archives), 
from whom he sought information and advice on specimens he collected and 
for whom he collected specimens. McFarlin’s early collections were deposited 
with the herbarium at Syracuse University. Later collections involving about 
5,000 specimens were sent to the herbaria at the University of Michigan and 
the University of Florida (McFarlin 1935). Some of his specimens have been 
deposited in the U.S. National Arboretum (Gann et al. 2002) and other locations. 

In addition to vascular plants, McFarlin was also interested in mosses and 
liverworts. He at one time had prepared displays of some of his specimens in 
mahogany cases (M. Bryant, pers.comm.). He eventually donated his bryological 
collections to the University of Florida and to the National Museum of Canada 
in Ottawa (K. Perkins, pers. comm.). That collection supported a publication 
(McFarlin 1937) in which he concluded that the local moss population in Polk 
County, was “relatively depauperate of species.” McFarlin collected in south 


610 BRIT.ORG/SIDA 22(1) 


Florida as well and was noted as having collected the first specimens for a hand- 
ful of species in that region of Florida (Gann et al. 2002). 

McFarlin’s major botanical exploration, which resulted in the collection of 
the bulk of the specimens, occurred in 1930 and 1931 while a graduate student. 
McFarlin collected throughout Polk County, a large (5,300 sq. km.) county lo- 
cated in the central part of Florida. At the time he was collecting the area was 
still relatively undeveloped. He made repeated collecting trips to several loca- 
tions, including Marion Creek near Haines City, Lake Deer near Winter Haven 
and Kissengen Spring near the Peace River south of Bartow. 

McFarlin never received his doctorate, but the reasons are unclear. Accord- 
ing to the University of Michigan's Transcript and Certification Office, he was 
enrolled as a graduate student from 1930 to 1933 (M. Henderson, pers. comm.). 
In a September 1, 1935, letter to J.K. Small he wrote of having completed his 
dissertation and informed. Small he was ready to submit it to his doctoral com- 
mittee. He wrote on April 29, 1935, to E B. Mains, one of his professors at the 
university and was told that. McFarlin should first submit his thesis to John H. 
Ehlers, chairman of his committee. Yet there is no record of his dissertation’s 
ever having been received at the University of Michigan (E. Voss, pers. comm.). 
Although McFarlin’s university transcript is unavailable because of claimed 
privacy concerns, the Certification and Transcript Office agreed to examine the 
transcript and said it does not indicate why Mr. McFarlin did not complete his 
degree. (M. Henderson, pers. comm.). 

McFarlin’s 1935 dissertation, however, has survived. A copy of the original 
document was completely retyped by Margaret L. Gilbert, a biology professor at 
Florida Southern College in Lakeland, Florida, in 1961, with duplicates given to 
several individuals and institutions within the state. The thesis is a tightly writ- 
ten work of 228 pages (as retyped), the bulk consisting of an enumeration of the 
vascular flora arranged by family. Each species is reported by scientific name, 
synonyms, and at times common names, with a brief statement of habitat and 
location. Nearly all entries also carry McFarlin’s field collections numbers for the 
species. A key is given to the species of genera with more thana single species. 

Asa supplement to the main body of the thesis, McFarlin provided a sta- 
tistical tabulation of its contents: 159 families, 560 genera, and 1131 species. He 
then recorded 131 species he regarded as weeds, and 53 species as introductions 
He gave further complete tabulations for the species of “high pine land,” “scrub,” 
“high hammock,” “flatwoods,” “prairie,” “low hammock,” “swamp,” “bayhead,” 
“cypress head,” “marsh,” “lakes,” “streams,” “clay pits,” “bogs,” “shallow ponds and 
drainage ditches,” “shores,” and “sandy beaches.” He followed this with lists of 
“epiphytes,” “parasites,” 11 and “saprophytes.” 

McFarlin recorded in the last pages of his thesis what he believed to be 2 
species new to the United States, 7 new to Florida, 7 species new to science, 7 
new varieties, 14 new forms, and 3 new combinations 


bya s3 


boas 


PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 611 


McFarlin’s 1935 thesis and his supporting specimens (mostly those at the 
University of Florida) have provided clues for further botanical exploration. His 
work, though unpublished and little known, offers a rare glimpse of the former 
distribution and abundance of many species that are now much less common 
in central Florida. 

A few examples are worth mentioning. McFarlin reported collecting the 
first U.S. specimen of Hairlike Spikerush (Eleocharis nana),a species that is still 
considered rare and has been collected from only two other Florida counties 
(Wunderlin & Hansen 2000). However, the collection of this specimen under- 
lines one of the problems with reviewing McFarlin’s work. The reported collec- 
tion site (Lake Lynch near Lake Alfred) is unknown and appears neither on 
modern maps nor on maps from that period, such as the 1927 Polk County Soil 
Survey that he cited in his dissertation. Several other sites where McFarlin re- 
ported collecting regularly, such as Highland Gully near Lakeland and Faulkner 
Hammock near Bartow, are similarly undocumented as to their locations other 
than being near some present-day city. In an undated letter to E.B. Maines, Mc- 
Farlin refers toa map “showing all of my collecting locations,” but no such map 
was included in any of the known copies of his doctoral dissertation. 

In some cases, McFarlin found species beyond their known ranges in Florida 
at the time, suchas the terrestrial orchid Wild Coco (Eulophia alta). Until then 
the species was known only from the Big Cypress Swamp area, according to 
Small (1933). 

McFarlin claimed (McFarlin 1935) to have collected a number of specimens 
that either represented species previously unknown to Florida or entirely new 
species. In the case of the new species for Florida, it appears possible that some 
of the claimed discoveries, such as Nash’s blue-eyed grass (Sisyrinchium 
brownei) or Ludwigiana brevipes, may have been misidentified since modern 
summaries of Florida’s botanical life, notably Wunderlin and Hansen (2000), 
do not include them. However, others suchas Verbena bonariensis and Commelina 
benghalis are known to exist in Florida and are actually quite common. 

McFarlin’s only published new species was a small- leaved sand holly that 
he named Ilex pygmaea (McFarlin 1932). He was unaware that the tree had been 
named previously, by Small in 1924, and his name is now considered a syn- 
onym of the scrub holly, [lex opaca var. arenicola. All but one of the six other 
new species mentioned in his unpublished thesis had no nomenclatural stand- 
ing and were largely overlooked. The exception, which has recently received 
attention, is the pink-flowered Scrub (McFarlin’s) Lupine, which McFarlin 
named Lupinus aridorum. This plant was collected by other workers as early as 
1900 (U.S. Fish and Wildlife Service 1993) but McFarlin was the first to con- 
sider it a separate species. McFarlin did not describe the species. It was not de- 
scribed until years after his death by Beckner (1982), who also knew the plant 
in the field, recognized its distinct features, and gave it proper treatment. Though 


612 BRIT.ORG/SIDA 22(1) 


related to Lupinus westidanus, a similar species of the Florida panhandle, it is 
quite separate in its range and is clearly an endemic of the central peninsula. 
McFarlin described the plant as “infrequent to locally abundant,” but with de- 
velopment of the area it has become quite rare, and is now classified as endan- 
gered by the state and federal agencies (Coile & Garland 2003). Asa result, the 
future survival of this species remains a concern. The only protected popula- 
tions of the plant occur at a federal preserve in Eagle Lake south of Winter Ha- 
ven and at a county park in the Orlando area (Kane 2003). Unprotected popu- 
lations persist in a handful of other sites in Central Florida. Establishment of 
other protected populations—including one site near Winter Haven where Mc- 
Farlin did much of his collecting—has been proposed (Kane 2003) and would 
seem toa fitting tribute to McFarlin’s work. 

McFarlin commented on other endemic plants of the central ridge. Another 
species he believed to be new was a small-leaved variety of grape that is wide- 
spread in the scrub habitat. He gave it the name Muscadina pygmea. Later ob- 
servers have noted the plant, but have never given it formal status by publish- 
ing its name and description. Recently McFarlin’s original specimens have been 
tracked down (in the Smithsonian Institution) and a paper is in press to give it 
formal recognition as a variety of the muscadine grape, Vitis rotundifolia (D.B. 
Ward, pers. comm.). 

McFarlin made interesting observations on the distribution of known en- 
demic species in the scrub habitat in Polk and adjacent counties, relatively little 
of which remains (Christman 1988). McFarlin’s comments were prescient re- 
garding this trend. For instance, he noted that the presence of Turkey Oak (Quercus 
laevis) was a general indication of land that was suitable for citrus, which is the 
crop that most frequently displaced scrub and sandhill habitat. Comparing his 
comments (McFarlin 1935) with those of present day observers helps us to un- 
derstand how dramatic that change has been. McFarlin described the beargrass, 
Nolina brittonidna, as “our most common species inhabiting the scrub and in- 
land sand dunes,” the crucifer, Warea amplexifolia, as “locally abundant,” the 
scrub plum, Prunus geniculata as “widespread,” the morning-glory, Bonamia 
grandiflora as “frequent in the sandy soil of the scrub,” and the milkweed, “As- 
clepids curtissii, as “frequent in scrub.” Now they are rare and classified as en- 
dangered or threatened (Coile & Garland 2003; Wunderlin & Hansen 2000). 

He left another legacy that was probably unintended. Although McFarlin 
recognized the problems of invasive exotic species—his manuscript mentions 
Ceasar Weed (Urena lobata), Australian Pine (Casuarina esquisetifolia) and oth- 
ers—he was responsible for some local introductions of other invasive plant 
species. Through his years in the nursery trade, McFarlin was responsible for 
the introduction of of the orchid tree, Bauhinia variegata, into Winter Haven 
(Burr 1974). This tree is classified as a Class | invasive species (Florida Exotic 
Pest Plant Council 2005). While operating the botanical garden at Highlands 


— 


PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 613 


] © LA] 


Hammock, McFarlin was apy ly for the introduction of the Flax 
Lily (Dianella caerulea), which continues to be a management problem in the 
park today (A. Hine, pers comm.). 


ACKNOWLEDGMENTS 


Ithank Daniel Ward and John Beckner for their encouragement and for provid- 
ing helpful comments during the preparation of earlier drafts of this account 
and Ward and Ron Gilmour for their review of the final draft. lalsothank Nancy 
Bissett for introducing me to Mr.McFarlin’s work in Polk County, Rita Lassiter 
for sharing her copies of J.K. Small’s correspondence, Edward Voss for help in 
locating relevant correspondence from the University of Michigan and Mary 
Brigham Bryant for sharing her childhood recollections. 


REFERENCES 


Autvarer, A.C. 1966. Highlands Hammock. Sebring Historical Society, Sebring, i 

Anonymous. 1969. James Brigham McFarlin. Obituary. Bradenton Herald, March 10, 1969. 

Becker, J. 1982. Lupinus aridorum J.B.McFarlin ex Beckner (Fabaceae), A new species from 
Central Florida, Phytologia 50:209-211. 

Burr, J.G. 1974. History of Winter Haven, Florida. Larry Burr Printing Co., Winter Haven, FL. 

CHRISTMAN, S. 1988.Endemism and Florida’s Interior Sand Pine Scrub. Final Project Report, 
Florida Game and Freshwater Fish Commission, Tallahassee. 

City Directory, WINTER Haven. 1928. R.L Polk Co. Jacksonville, FL. 

Cote, N.C. and M.A. Gartanp. 2003. Notes on Florida’s endangered and threatened olants. 
Florida Dept. of Agric. and Consumer Services, Gainesville. 

Fioriva Exoric Pest Plant Councit. 2005. Invasive plant list. http://www.fleppc.org/list/list.hAtm. 
Tallahassee, FL. 

Gann, G.D., K.A. Braptey, and S.W. Woopmansee. 2002. Rare plants of South Florida: Their his- 
tory, conservation, and restoration. Institute for Regional Conservation. Miami, FL. 

Kane, S.2003. Historical and current distributions and population status of Lupinus aridorum 
McFarlin ex Beckner. Masters thesis, University of Central Florida, Orlando. 

McFarun, J.B. 1932. Two new evergreen holly from Central Florida. Rhodora 34:16-18. 

McFaruin, J.B. 1935. Flora of the central portion of the Lake Region District of Florida. 

McFaauin, J.B. 1937. Mosses of Polk County, Florida. Bryologist 40:49-5/7. 

McFaauin, J.B. 1939. Highlands Hammocks Trails. A guide to the plants along the Laurel 
Oak Trail. Highlands Hammock State Park, FL. 

Smait, J.K. 1933. Manual of the southeastern flora. Published by the author. New York. 

U.S. FisH AND WiLouiFe Service. 1993. Endangered and threatened status for seven Central 
Florida Plants. Fed. Reg. 58(79):25746-25755. 

WUNDERLIN, R.P.and B.F. Hansen. 2000. Flora of Florida.Vol.1.Pteridophytes and gymnosperms. 
Univ. Press of Florida, Gainesville. 

WUuNDERLIN, R.P. and B. HANseN. 2003. Vascular plants of Florida, 2nd Edition. Univ. Press of 
Floria, Gainesville. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


WILLIAMS. Justice, C. Rircuié Bett,and ANNE H. Linpsry. 2005. Wild Flowers of North 
Carolina (Second Edition). (ISBN 0-8078-5597-9, pbk.). University of North 
Carolina Press, P.O. Box 2288, Chapel Hill, NC 27515-2288, U.S.A. (Orders: 
www.uncpress.unc.edu/books/T-7662.html). $29.95, 376 pp., 553 color and 
6 b&w illus., 6 tables, | chart, appendices, bibliography, index, 5 1/2" x 
ol/2". 


Edition 2 supplants the original from 1968 by Justice and Bellit includes 100 more species than the 
With conservation and preserva- 


original and provides brief notes on toxicity and medicinal uses. 
‘irst in mind, the authors note that “Our goal in this presentation of 500 native or naturalized 


tion 
plants in North Carolina is to open eyes, minds and hearts to the story of the state’s wild flowers, 
their beauty, their interesting attributes, their uses, and, in many cases, their plight.” Most pages in- 
clude a photo and notes for each of two species, and Appendix | gives an outline of horticultural 
information for each species. The Edition 2 photos are better reproduced, often sharper, sometimes 
cropped differently, but usually reduced in size up to 30 percent. For Edition 3, better planning of 
increase image sizes and decrease the large areas of white space on so many of 
Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102- 


layout surely could 


the pages. Guy Nesom, Botanical 


4068, U.S.A, 


SIDA 22(1): 614. 2006 


FLORISTIC DIVERSITY AND COMPOSITION 
OF TERRA FIRME AND SEASONALLY INUNDATED PALM 
SWAMP FORESTS IN THE PALMA REAL WATERSHED IN 
LOWER MADRE DE DIOS, PERU 


Fernando H. Cornejo Valverde John P Janovec 
Botanical iesedien Institute of Texas — Peru Botanical Research Institute of Texas 
SN a Ae 09 Pecan St. 
cae de Dios, PERU Fort Worth, Texas 102-4060, U.S.A. 
fcornejo@brit.org jjanovec@brit.org 


Mathias W.Tobler 


Botanical Research Institute of Texas 
09 Pecan St. 


Fort oe si - 102-4060, U.S.A. 
tobler@gmx.net 


ABSTRACT 


We report the results of a tree species inventory of seven hectares of lowland subtropical moist forest 
in the Palma Real River watershed of lower Madre de Dios, Peru, where we discovered a type of palm 
swamp forest vegetation that has not previously been studied. The goal of the study was to investi- 
gate the forest vegetation of the Palma Real watershed with special emphasis on a comparison be- 
tween the wetland forest vegetation and the surrounding terra firme forests. A total of 3534 trees 
>10 cm DBH belonging to 442 species in 62 families were measured and identified in the seven 
plots. Three distinct forest types resulted from Principal Components Analysis (PCA) and WPGMA 
cluster analysis of both family and species abundance data, including two types of terra firme for- 
ests and a palm swamp forest. The five terra firme forest plots sampled, with a range of 104-138 
species, were less diverse than other one-hectare forest plots reported from the region, which have a 
diversity ranging from 151-197 species. The two palm swamp forest eee with 126 and 167 tree spe- 

cies, were more diverse than surrounding terra firme forest plots sampled in this study. One of the 
palm swamp forest plots was extremely diverse with 167 species, ae seven species less than the 
average diversity across 14 one-hectare terra firme plots sampled 2 other investigators in Madre de 
Dios, Peru. Both palm swamp forest plots had a greater number of stems distributed across smaller 
size classes. Both Fabaceae and Arecaceae were ane most important plant families across all plots. 

The Arecaceae contributed significantly to g the palm swamp forest plots, composing 
16-25% of all stems compared to only 9% of the stems in terra firme plots of the region. This high 

abundance of the palm family is from where we derive the name we are applying to this wetland 
forest type in the Palma Real watershed. Using satellite imagery we illustrate the importance of the 
extent of these palm swamp forests throughout the region 


RESUMEN 


Reportamos los resultados de un inventario floristico de siete Becuce en un pees SHE ADICH 


amedo en la cuenca de la quebrada Palma Real, en Madre de Dios, Per 
fue investigar la vegetacion de la Quebrada Palma Real con especial Hee ae en una comparacion 


SIDA 22(1): 615 — 633. 2006 


616 BRIT.ORG/SIDA 22(1) 


entre un tipo de vegetacion de Si antano con pn panier y la uegetacion de terra lirme circundante. i n 
4 AyD 


total de 3534 arboles mayoresa 
fueron medidos e identificados en siete parcelas. Tres tipos de bosque résultaron de los analisis de 
componentes principales y analisis de agrupamiento WPGMA mediante datos de abundancia por 


familia y por especie en los dos tipos de ee aoe Las cinco dices yess en terra a me, 


con un rango de 104-138 especies por parcela, fuer sas que otras p 

en la region, las cuales tienen un rango de 151-197 especies. Las dos parcelas cima en . 
pantanoso de palmeras, con 126 y 167 especies, fueron mas diversas que las parcelas de terra firme 
circundantes. Una de las parcelas del bosque pantanoso de palmeras fue extremadamente diversa 
con 167 especies, solo siete especies menos que el promedio reportado para |4 otras parcelas estudiadas 
por otros investigadores en Madre de Dios. Ambas parcelas del bosque pantanoso con palmeras 
tuvieron una alta censidad de tallos sania en ~ cl ases diamémetricas mas pequenas, lo cual 
podria ser ion de laalt i . Fabaceae y Arecaceae fueron las familias 


mas pee para todas las eos orn contribuy6 significativamente a distinguir el 
bosque pantanoso con palmeras, compre aie coen te el 16-25% Ge i ee ss uaseuiak con 


fs a lt 


solo el 9% de tallosen | DOs a de Cerra firme ade la region FE 


el nombre que ste tipo de bosque muy humedo en la cuenca de la Quebrada de 


Palma Real. Usando imerenee csielicales nosotros ilustramos la importancia por su extension, de 


estos bosques pantanosos de palmeras en la region. Finalmente nosotros instamos a Mowe mas 


estudios de largo plazo sobre - eee, pees en a region, con é ome en la importancia de 


estos bosques de Palma Real e trac : i, Lecythidaceae) y otro: 
recursos para la region y sus ee 
INTRODUCTION 

A large forest plot network has accumulated in the last decade from the Ama- 
zonian region. The highest tree species diversity has been reported from the 
western Amazon where Gentry (1988) and Valencia et al. (1994) recorded 283 
and 307 species respectively in one-hectare forest plots. Oliveira and Mori (1999) 
found 285 species per hectare on very poor soils and under reduced rainfall in 
central Amazonia, which is essentially the same species diversity as found by 
Gentry in western Amazonia. In an analysis of data from 97 existing one-hect- 
are forest plots from the Amazon, including the Guiana Shield, Ter Steege et al. 
(2000) showed that in all regions terra firme forests harbor a greater number of 
tree species per unit than any other forest types. Patterns of high tree species 
diversity have been attributed to high rainfall and more nutrient-rich soils of 
the western Amazon (Gentry 1988). 

Nebel et al. 2001) and Kvist and Nebel (2001) emphasize the importance 
of wetland forests to both the ecological diversity and resource use in the Peru- 
vian Amazon. While many forest plot inventories in the Amazon have focused 
on the diversity of terra firme and floodplain forests, few studies in the region 
have dealt with wetland vegetation. For example, only two swamp forest plots 
from the Amazon were available for inclusion in the large studies by Ter Steege 
et al. (2000, 2003), but those were excluded from the final analyses and spatial 
modeling. There is clearly a lack of data from wetland habitats in the Amazon. 
This minimal coverage of wetland vegetation leads toa generalized view of the 


CORNEJO ET AL., FLORISTIC PERU 617 


— 


complexity and diversity of these ecosystems, as discussed by Kalliola et a 
(1991), who argue for more focused studies. 

The Tambopata-Candamo region and Manu National Park are two well- 
studied protected areas of Madre de Dios, Peru, that hold world records in diver- 
sity of various organisms, such as birds, insects, and mammals. National re- 
serves, such as Tambopata, allow for protection and sustained use of plant 
resources. The Palma Real watershed is inside the Tambopata National Reserve 
and the Bahuaje-Sonene National Park, where management plans are needed 
since plant and animal extraction is carried out by indigenous and local people. 
Yet, no floristic treatment exists for the region and only a few small areas are 
represented by checklists. 

In the Palma Real watershed and the surrounding region, there exists a vast 
area of terra firme forest and patches of seasonally inundated swamp forests. 
This mosaic of forests extends from the Tambopata Reserved Zone of lower 
Madre de Dios, Peru, to the Pampas del Heath in the border region of Peru and 
Bolivia. The seasonally inundated forests of this region are characterized by a 
distinctive topography that consists of small raised islands separated by shal- 

ow water channels during the rainy season or heavy rains in the dry season, 
and they sit upon a deep, fine, heavy gray soil. Because of the abundance of 
palm species in our study area we refer to this wetland vegetation as palm 
swamp forest. We have reviewed available literature that relates to the diversity, 
ecology, and distribution of wetland vegetation in the area, and we have found 
no existing description of this specific wetland vegetation. The palm swamp 
forests of Palma Real are different in species and family diversity and composi- 
tion than the palm swamp forest communities of the northern Peruvian Ama- 
zon described by Kalliola et al. (1991). Although Haase (1990) described palm 
swamp communities in northern Bolivia, there is no description that matches 
the palm swamp forests of the Palma Real watershed. Nor are these palm swamp 
forest communities covered by Beck (1984). The goal of this study was to inves- 
tigate the forest vegetation of the Palma Real watershed in lower Madre de Dios, 
Peru, with special emphasis on a comparison between these palm swamp for- 
ests and the surrounding terra firme forest vegetation. 


METHODS 


Study Site 

The study was conducted in the Palma Real watershed, a tributary of the Madre 
de Dios River, approximately 28 km to the southeast of Puerto Maldonado, the 
capital and the principal city of the Department of Madre de Dios, Peru (Fig. 1). 
Nested within the Tambopata-Candamo National Reserve and the Bahuaje- 
Sonene National Park, this area of lowland subtropical moist tropical forest (Fos- 
ter et al. 1994) receives about 2208 mm of precipitation annually and the aver- 
age temperature is 24°C (mean over 36 years). During the driest months, May to 


BRIT.ORG/SIDA 22(1) 


618 
YP 
9 
Naf q C7 
oe Rivet 
ae DY” - 
gate 
= M QS 
ZY, ¢ 
Kn rd ) Limon { 
° ©) 
< Puerto Maldonado 
* fro A : 
bd C & Oculto 
. x 
yw? & eo 
fs > 
i QW Ka) e 
) < 
Q& Lf @ 
wS NY S 3s Settlements 
ARN) La Viuda A Plots 
G, y ‘Soh ~ N 
; km 
. 0 5 10 #15 
Pd 
Fic. 1. Location of the three si here f pl li based, Limon, Oculto, and La Viuda. 
hich 


September, the area receives occasional cold winds from the antarctic, w 
decreases daily temperatures to a minimum of 7-8°C, although this phenom- 
enon of cold fronts, known as “friajes”, lasts for only a few days at a time. 

The general topography is flat, dissected by small streams, most of which 
are without water during the dry season. Three main types of habitats exist in 
the Palma Real watershed, forming an interesting mosaic of vegetation zones: 
(1) high terrace non-inundated forest (terra firme); (2) palm swamp forests - 
areas with poor drainage and seasonal inundation causing the formation of 
swamp-like forests; and (3) seasonally inundated floodplain forest. This study 
focused on terra firme and palm swamp forests, which are the most extensive 
forest types in the study area. The Brazil nut tree (Bertholletia excelsa, 
Lecythidaceae) is abundant in large areas of the terra firme forests in this re- 
gion but it is absent from others. Seeds of the Brazil nut tree are extracted from 
the Palma Real watershed by harvesters (castaneros) every year during Janu- 
ary-March, making this an economically important region in Madre de Dios, 
Peru. The forests of the Palma Real watershed maintain a healthy fauna with 
jaguars, ocelots, tapirs, peccaries, deer, and at least six species of monkeys. No 


J 
logging activities have been conducted in this region since twenty years ago 


Ss 


CORNEJO ET AL., FLORISTIC DIVERSITY IN LOWER MADRE DE DIOS, PERU 619 


when many Spanish Cedar trees (Cedrela odorata, Meliaceae) were extracted. 
Then terra firme forests of Palma Real that are cl ized by high Brazil nut 
tree density have been divided by the Peruvian government into forest conces- 
sions of approximately 1000 hectares each and given to local extractors living 
in Puerto Maldonado. 

Soils.—The soils of the terra firme forests are dominated by red clay and 
those of palm swamp forests by gray clay. Within the Palma Real zone, three 
representative sites were selected for quantitative studies of floristic diversity 
and composition (Figs. 1, 6): (1) Limon (12°32'19"S; 68°51'40"O), close to the Madre 
de Dios River; (2) Oculto (12°39'03'S: 68°55'40"O), in the middle of the zone; and 
(3) La Viuda (12°47'43'S: 68°58'07"0), to the extreme south of the zone. 


Field Data 

Seven permanent one-hectare forest plots were established in the Palma Real 
watershed, divided between Limon (3 plots), Oculto (3 plots) and La Viuda (1 
plot). Within each site, one plot was established in terra firme (TF) forest witha 
known high density of 5-7 individuals of Brazil nut trees per hectare (Limon 1, 
Oculto 1, La Viuda 1). One plot in each of Oculto and Lim6n was established in 
TF forest with known absence of Brazil nut trees (Oculto 2 and Limon 2). In 
Oculto and Limon one plot was also established in palm swamp (PS) forest 
(Oculto 3 and Limon 3). 

Each plot consisted of a square area of 100 x 100 m, divided into 25 sub- 
plots of 20 x 20 m. Each corner of the one-hectare forest plot is marked witha 
PVCstake labeled with the respective X,Y coordinate in red paint. All individu- 
als, including palms, 10 cm or greater in diameter at breast height (DBH) were 
measured, mapped and identified to family, genus, and species or morphospecies 
when definite species determination was not possible. All individuals with 
multiple stems were considered as one individual, and the diameters of each 
stem were summed for basal area calculations. 

Every species is represented by duplicate voucher specimen collections of 
one or more individuals. All identifications were made in the La Molina Her- 
barium (MOL )at the Agraria University in Lima, Peru, where duplicate voucher 
specimens were deposited. The Cronquist system of classification of flowering 
plants was used for the purposes of this project. 


Data Analysis 

Relative diversity, relative density, and relative dominance were calculated for 
each plot. The Importance Value Index (IVI) was calculated at the species and 
family level for each plot using the sum of the three measures, relative diversity, 
relative density, and relative dominance. Sorensen Similarity Index (Serensen 
1948) was calculated for all species found in the plots and for the 20 most abun- 
dant species. The Sorensen index is 2 * S}2 / (S1 + S2), where S)2 is the number of 
species common to both sites and S;and $2 are the total number of species found 


620 BRIT.ORG/SIDA 22(1) 


at site one and two. Plots were also grouped based on species and family abun- 
dance using Principal Components Analysis (PCA) and WPGMA cluster analy- 
sis (MVSP 3.12, Kovach Computing Services). 

Comparing the raw number of species between plots without accounting 
for differences in stem density can give misleading results. To avoid this prob- 
lem we used individual based rarefaction curves as suggested by Gotelli and 
Colwell 2001). Rarefaction curves were calculated using the software package 
EstimateS 7.5 (Colwell 2005). 


RESULTS 


Forest Structure 

A total of 3534 trees =10 cm DBH belonging to 442 species in 62 families were 
measured and identified in the seven plots (Table 1). Three distinct forest types 
resulted from PCA: (1) palm swamp forest (Limon 3 and Oculto 3); (2) Limon 
terra firme forest (Limon | and Limon 2): and (3) Oculto/La Viuda terra firme 
forest (Oculto 1, Oculto 2 and La Viuda) (Figs. 2, 3). PCA resulted in almost no 
difference between the species (Fig. 2) and family level (Fig. 3),a pattern previ- 
ously suggested by Terborgh and Andersen (1998). 

Table | shows the principle structural characteristics for each plot. The palm 
swamp (PS) forests (PS) in both Limon and Oculto have a higher number of 
species than the terra firme (TF) forests at the same sites. Limon 3 has the high- 
est number of individuals and species of all the plots while the two TF forest 
plots at Oculto are the least diverse. The rarefaction curves (Fig. +) confirm these 
results and show that the higher diversity of the PS forest plots are not simply 
due toa higher stem density. The PS plots have a lower basal area than the TF 
plots (215 vs. 27.3). Distribution of trunk diameter classes shows the inverse J- 
shaped curve, witha mean proportion of 69% of individual trees found between 
10 and 20 cm DBH. 


= 


Composition 

Monimiaceae was the most abundant family in the TF forest plots, followed by 
Arecaceae and Fabaceae (Table 2). In PS forest plots Arecaceae and Fabaceae 
were the most abundant families. Looking at the 20 most abundant families 
(Table 2), Lauraceae, Moraceae, Violaceae, Lecythidaceae, Chrysobalanaceae, 
Sapotaceae, Burseraceae, and Cecropiaceae were found in both TF and PS for- 
ests, whereas Myristicaceae, Strelitziaceae, Linaceae, Bixaceae, Rubiaceae and 
Flacourtiaceae were found only in TF forests. Of the 20 most abundant families, 
Myrtaceae, Dichapetalaceae, Anacardiaceae, Clusiaceae, Ochnaceae, Sterculiaceae, 
and Combretaceae were restricted to the PS forests. The ten most abundant fami- 
lies represented 64% of all individual trees found. The most species-rich family 
across all plots was Fabaceae, followed by Lauraceae, Moraceae, Sapotaceae, 
Chrysobalanaceae, and Annonaceae. The ten richest families represented 54% 


CORNEJO ET AL., FLORISTIC DIVERSITY IN LOWER MADRE DE DIOS, 


PERU 


Taste 1. Structural characteristics from 1 hectare plot. 


Limén1  Limén2 LaViuda Oculto1 Oculto2 Oculto3 Limén3 
No. of stems 499 Sop 473 422 508 492 608 
No. of species 138 130 130 109 104 126 167 
Basal area ( m7’) 32.23 23.77 25.23 29.25 26.22 21.86 21.07 
Categories by diameter (%) 
10-20 cm 3.4 70.5 69.5 64.9 65.9 74.6 74.5 
20.1-40 cm 27.6 23.5 22.2 23.0 24.8 203 21.9 
40.1-80 cm 74 5.1 7.0 10.2 8.3 4.3 35 
> 80cm 1.6 0.9 1.3 1.9 1.0 0.8 0.2 
No. of species in 50% IVI 16 16 18 13 13 14 20 
No. of families in 50% IVI 39 42 4] 4] 40 42 45 
4.09 —- 
3.27 —— 
2.45 —— 
La Viuda 
A 
Oculto 3 
A Oculto 1 
Oculto 2 0.82 -- 
£ | —+—_-__ SEE EEEEEEEE eee 
a | 
Ne) -3.27 -2.45 -1.63 -0.82 0.82 1.63 2.45 3.27 4.09 
i672 7. 
Limén 3 
-1.63 — 
Limon 2, ~2:45-- 
A 
Limén 1 
A» = 
Axis 1 


Fic. 2. Ordinati f inci 
r t a 


622 BRIT.ORG/SIDA 22(1) 


Limon 1 26.7—-+- 


A 
Limon 2 


c SIXW 


-6.7 + 
La Viuda 
A A 


A 
Oculto 1 aan Oculto 3 


A 
Oculto 2 -20.9—- 


Nedinati £4 Deine 
Fic. 3. Ordination of a | p f y J 


of all species. Table 3 summarizes the Importance Value Index (IVD for the 20 
most abundant families per plot, per forest type, and across all plots, emphasiz- 
ing the importance of Arecaceae in PS forest plots, and the Fabaceae across all 
forest plots. 

Table 4 lists the twenty most abundant species in each of the seven I-ha 
forest plots. There were only seven species that occurred in all seven plots. Twenty 
species were found to occur in all five TF forest plots while 62 species were found 
to occur in both of the PS forest plots. A total of 243 species were found to occur 
in only one plot out of the seven. Out of the total of +42 species found across all 
the plots, 124 occurred only in the PS forest and 212 only in the TF forest. A 
total of 106 species were found to occur in both forest types. About +0% of the 
species were represented by only one individual and 65% by | to 3 individuals. 
Similarity 
Table 5 shows the Sorensen Similarity Index for all plots. Values range from 0.25 
to 0.55 for all species and from 0.10 to 0.75 when only the 20 most abundant 


CORNEJO ET AL., FLORISTIC PERU 623 
180 9 
160 74 a MSc. 
Limon 1 i 
La Viuda Lim6én 3 
140 5 
120 4 Te Limon 2 
Oculto 3 
w 1007 
= 
) 
a Oculto 2 
® 894 Oculto 1 
60 4 
40 4 
20 4 
Q T T ; T T T 1 
0 400 200 300 400 500 600 700 
Individuals 
D £. + L I L L + 1 £ j j ! J L ££ 
Fic.4 g I p 


species were included. The 20 most abundant species, on average, represent 60% 
of all individuals (Range: 54-69%). Similarity analysis shows the same patterns 
already observed in the PCA. First, plots within the same forest type (TF or PS) 
are more similar to each other. Considering only the 20 most abundant species 
in each plot, this pattern becomes even more apparent. The TF forest plots from 
the same site (e.g., Oculto | and 2 or Limon 1 and 2) share a higher number of 
species with each other than they do with plots from different sites. Figure 5 
provides the results of WPGMA cluster analysis based on species abundance, 
which shows that PS forest plots cluster together, as do TF forest plots from the 
same sites. When all species are considered, PS forests are more similar to the 
TF forest plots at the same site than to those farther away. However, this is not 
true when we consider only the 20 most abundant species. In this case, Limon 
3 shares the same number of species with Limon Las it does with Oculto 2, and 
Oculto 3 and Oculto | are the least similar plots. 


DISCUSSION 
The Palma Real watershed presents a unique seasonally inundated PS forest 
habitat that is very different from the TF forests of the region. Based on species 
abundance, the two PS forest plots are clearly distinguished from the TF forest 
plots as shown by results of both PCA (Figs. 2, 3) and WPGMA cluster analysis 
(Fig. 5), as well as similarity index analysis (Table 5). Euterpe precatoria is the 


624 BRIT.ORG/SIDA 22(1) 


nH haldt rh +f 


Taste 2. The 20 most diverse families in the two main forest types. Families i those tnat 
are among the top 20 in both forest types. Number of individuals are average by hectare 


Terra firme Forest palm swamp Forest 
Family No. of No. of Species Family No. of No. of Species 
individuals individuals 
ceae 41.6 39 Fabaceae 78 35 
Lauraceae 30.6 26 Annonaceae 12 12 
26.8 22 Chrysobalanaceae 18 12 
Sapotaceae 9.6 1/7 Lauraceae 75 1] 
Chrysobalanaceae 7.6 14 Sapotaceae 16 11 
Meliaceae 12.4 12 Moraceae 33.5 10 
Annonaceae 94 1] Burseraceae 15.5 8 
Cecropiaceae 20.2 10 Arecaceae 128.5 7 
Monimiaceae 53 9 Apocynaceae 10.5 7 
Flacourtaceae 6 9 Anacardiaceae 11.5 6 
pocynaceae 78 7 Lecythidaceae 24 7 
Ar ea 52.8 6 Clusiacea 10.5 4 
Burseraceae 19.8 6 tacea 15 4 
Myristicaceae 36.8 5 Cecropiaceae Ves) 3 
Rubiaceae 7.2 4 M ea 8 3 
Lecythidaceae 124 4 Dichapetalaceae 11.5 2 
Violaceae 26.6 2 Violaceae 28.5 2 
Linaceae 9.4 2 Ochnaceae 10.5 2 
Strelitziaceae 14.2 | Combretaceae 8 2 
Bixaceae 9 | Sterculiaceae 10 2 


most abundant species in PS forest plots, Limon 3 and Oculto 3, contributing 
significantly to the major differentiation between the PS and TF forest plots. 
Astrocaryum murumuru, Attalea butyracea, Brosimum laevis, Chimarrhis sp. 1, 
Dipteryx micrantha, Eriotheca globosa, Manilkara bidentata, Ouratea weber- 
baueri, Rheedia sp. 2, Terminalia oblonga, and Triplaris americanaare restricted 
to PS forest plots. Calyptranthes bipenis, Iryanthera juruensis, Lauraceae sp. 1, 
Lauraceae sp. 2, Pourouma minor, and Rubiaceae sp. 2 are restricted to TF forest 
plots. Based on our results, there are two groups of TF forest plots as demon- 
strated by Figures 2, 3, and 5. The first group of TF forest plots, Limon | and 
Limon 2, is distinguished primarily by the abundance of Brosimum dlicastrum, 
Pseudolmedia laevigata, and Rinoreocarpus ulei, and secondarily by 
Phenakospermum guyanense. The second group of TF forest plots, Oculto 1, 
Oculto 2, and La Viuda, is distinguished by the abundance of Minquartia 
guidnensis, Oenocarpus batua, Roucheria punctata, Roucheria sp. 1, Rubiaceae 
sp. l, Socrotea exorrhiza, and Tontolea corymbosa. 

Analysis of family abundance demonstrates similar results (Fig. 3, Table 
3). The families Fabaceae and Arecaceae are the most abundant families across 


~— 


CORNEJO ET AL., FLORISTIC PERU 625 


Taste 3. Family Importance Value (FIV) of the 20 most abundant families for each plot, both forest 
and across all plots (sorted by Total column). The FIV was calculated as the sum of relative 
a relative density, and relative dominance. 


Family LaViudal Limén1 Limon2 Limén3 Ocultol Oculto2 Oculto3 PS TF Total 
Fabaceae 35.39 33.25 42.05 52.04 25.96 29.65 47.27 48.96 33.55 37.28 
Arecaceae 16.76 1446 17.85 31.91 1865 2238 48.26 3836 16.34 22.32 
Moraceae 13.14 3699 39.55 2583 918 3.01 27.71 27.37 2048 21.30 
Lecythidaceae 2045 3071 490 17.16 3798 1741 949 1345 22.05 20.11 
Lauraceae 22.78 21.32 1584 520 23.29 2639 684 695 2239 19.16 
Monimiaceae 1735 1340 19.64 1.62 2063 16.20 426 325 1765 13.28 
Sapotaceae 9.24 1275 745 1436 800 10.78 7.61 11.59 10.56 11.31 
Burseraceae 6.26 8.81 10.92 11.01 1286 15.82 453 810 1036 9.52 
Myristicaceae 1242 969 669 446 1280 2457 141 3.13 12.01 9.17 
Cecropiaceae 8.40 11.79 16.22 362 931 17.03 425 3.51 11.63 9.07 
Vidas 5.80 5.51 606 1040 9.63 573 1452 12.06 7.26 8.69 
Violac 2.58 22.93 12.96 1398 3.11 240 448 940 859 852 
eee 8.25 5.53 662 536 440 950 1222 848 696 7.31 


Flacourtiaceae 10.37 0.97 7.12 5.91 7.30 5.65 108 385 592 5.68 
Anacardiaceae 2.11 499 477 5.61 5.23 450 875 7.25 460 5,25 
Clusiaceae 6.18 3.7) 3.71 566 290 3.29 362 443 436 461 


all PS and TF forest plots, although they are more abundant in PS forest plots. 
The Lauraceae and Monimiaceae are more abundant in TF forest plots. The 
Violaceae and Moraceae are much more abundant in all the Limon plots in- 
cluding the PS forest plot Limon 3, distinguishing all the Limon plots from the 
other plots on axis lin the PCA (Fig. 3). The Dichapetalaceae and Polygonaceae 
are found only in PS forest plots, helping to distinguish them from TF forest 
plots. A higher abundance of the Bombacaceae in PS forest plots also helps to 
distinguish them from the TF forest plots. The first TF forest plot group of Limon 
1 and 2 is distinguished by a higher abundance of Lauraceae species than the 
rest of the plots, as well as by the Staphylaceae, which only occurs in Limon 1 
and 2 TF forest plots. The second TF forest plot group of Oculto l and 2 and La 
Viuda is distinguished by a high abundance of Myristicaceae, as well as by the 
Bixaceae, Hippocrateaceae, and Linaceae, which only occur in those plots. 

PS forest plots of Palma Real have a higher abundance of palms in com- 
parison to the TF forest plots, making the Arecaceae one of the most significant 
families distinguishing the different forest types (Table 2 & 3). This is especially 
apparent in Oculto 3, where palms account for the top three and sixth most 
abundant species sampled. Palms compose 25% of the stems in Oculto 3 and 
16% in Lim6n 3. In comparison, in all TF forest plots in the region palms make 
up only 9% of all stems. Euterpe precatoria, the most common palm in the area, 


= 


Tas.e 4. The twenty most abundant species in each of the seven one-hectare forest plots. 


LaViuda1 


Oculto1 


Oculto2 


Limon] 


Bixa arborea 

sic sp. | 

Siparuna decipien 
al chrysophylla 
Oenocarpus bataua 
Socrotea exorrhiza 
Euterpe precatoria 
Lauraceae sp. 1 
Roucheria punctata 
Iryanthera juruensis 
Tachigali polyphylla 
Calypthrantes bipenis 
Tetragastris a 
Iryanthera lae 
hoo sp. 1 
Phenakospermum guyanense 
Pourouma minor 
Rubiaceae sp. 2 

Inga gereauana 


Siparuna decipiens 
Siparuna $ 

Euterpe precatoria 
Iryanthera laevis 
Lauraceae sp. 1 
Ouratea sp. 1 
Oenocarpus bataua 
Protium 5 
Tetragastris panamensis 
Iryanthera juruensis 
Roucheria punctata 
Le polyphylla 


nt mb 
Tontolea corymbosa 
Hirtella giandistiotla 
Bertholletia excelsa 
Pourouma minor 
Trichilia quadrijuga 
Phenakospermum guyanense 
Hebepetalum humirifolium 


Iryanthera laevis 

Siparuna s 

Iryanthera juruensis 
uterpe precatoria 

toe oo 

Laurac sp. 1 

oe bataua 

Pourouma minor 

Eschweilera coriacea 

Phenakospermum guyanense 

Protium $s 

Socrotea exorrhiza 

Bixa arborea 

Siparuna decipiens 

Tetragastris panamensis 

Tontolea corymbosa 

Ouratea sp. 1 

Lacistema aggregatum 
Roucheria punctata 


NN TWO OO 


Leonia glycicarpa 
Rinoreocarpus ulei 
Euterpe precatoria 
Siparuna decipiens 
Iryanthera juruensis 
Lauraceae sp 
Pseudolmedia laevis 
Siparuna sp. 1 
Brosimum alicastrum 
Lauraceae sp. 2 

Guarea macrophylla 
Inga auristillae 
Lauraceae sp.9 
Pseudolmedia laevigata 
Socrotea exorrhiza 
ee panamensis 
Turpinia sp. 1 

Gall ulei 

Celtis schippii 


(2D) 


ON: (ONS) SS SST SN 


979 


(L)@Z VIS/9YO'LIYS 


Taste 4 (continued from left) 


Limon2 


Limon3 


Oculto3 


Euterpe precatoria 

Siparuna decipiens 
Phenakospermum guyanense 
Rinoreocarpus ulei 

Siparuna 

Tetragastis panamensis 


imum alicastrum 


pee nner 
p 


pouroura guianensis 
Lauraceae 

Pourouma minor 

Cecropia eae 
Eschweilera c 

Pseudolm ci ee 
Tachigali s 

eae vargasil 
Geissospermum reticulatum 


Euterpe 
Leonia glycica 

ee eens ence 
Eschweilera coriacea 
Tachigali chrysophylla 
Socrotea exorrhiza 
Rinoreocarpus ulei 
Ouratea cf. weberbaueri 
Tetragastris panamensis 


Brosimun lactescens 
Manilkara bidentata 
neuen tessmannil 
Ingae 

aay iis 
ees see 
Chimarrhis 


aude ese 


Rheedia 


CONN WWM 


Pl, SSS SSP Oe (COs AONE: (NO a 


Euterpe precatoria 
Socrotea exorrhiza 
Astrocaryum murumuru 
Tachigali chrysophylla 
Theobroma subincanum 
Attalea butyracea 
Tapura tessmannii 
Terminalia oblonga 
Eschweilera coriacea 
Leonia glycicarpa 
Brosimun lactescens 
Hirtella excelsa 

Triplaris americana 


Brosimun guianensis 
Cecropia sciadophylla 
Cordia cf. scabrifolia 
Inga gereauana 


JLSGO 1a TV La OfaNaQ) 


nud’ 


£79 


628 BRIT.ORG/SIDA 22(1) 


| 


Taste 5. Number of common species (above) and S@rensen Similarity In v) by one-hectare 
plot. The diagonal shows the total number of species for each plot. Alle species EM 20 most abundant 
species. 


La Viuda Limon 1 Lim6n 2 Limon 3 Oculto 1 Oculto 2 Oculto 3 
LaViuda 130 51/8 56/10 43/5 55/13 46/13 32/4 
Limon1 038/040 138 61/10 56/5 49/7 49/8 36/3 
Limon2 043/050 045/050 130 50/7 48/10 52/10 32/5 
Limon3 0.29/0.25 037/025 034/035 167 40/4 44/5 64/8 
Oculto1 046/065 040/035 040/050 0.29/0.20 109 59/15 34/2 
Oculto2 039/065 040/040 044/050 032/025 055/075 104 33/4 


Oculto3 0.25/0.20 0.27/0.15 0.25/0.25 044/040 0.29/0.10 0.29/0.20 126 


| al Oculto 3 
Limon 3 


Limon 2 


Limon 1 


Oculto 2 


Oculto 1 


La Viuda 


Fic 5 n A f WPGMA 1 | oe | 4 £. 1 7 . L 4 


is among the five most abundant species in six of the one-hectare forest plots 
and it is the most abundant species sampled in both of the PS forest plots, with 
88 stems in Lim6n 3 and 61 stems in Oculto 3. Socrotea exorrhiza is also an 
important species in PS forest plots, ranking as the sixth most abundant species 
in Lim6n 3 with 15 stems, and the second most abundant species in Oculto 3, 
with 32 stems. Astrocaryum murumuru is the third most abundant species in 
Oculto 3, with 31 stems. In Oculto 3, Attalea butyracea, known locally as 
“shebon’, is the sixth most abundant species, from which has come the local 
common name of this forest type, shebonal, but this is not true for Lim6n 3. 
One of the most important families in the TF forest plots was the 
Monimiaceae (Figs. 2, 3), making this the first report of high abundance and 
dominance of this family in any tropical forest plot study. This family was rep- 


CORNEJO ET AL., FLORISTIC PERU 629 


2] 
ry 


, Limén, Oculto, and La Viuda, and the 


HG ft bates im satin image from aay 2001 showing ine 


bw J 


the border of Peru and Bolivia. 


resented by two species of the genus Siparuna that were common in areas of 
tree fall gaps. The presence and abundance of Tachigali sp.,a monocarpic spe- 
cies of the Palma Real watershed, might explain the presence of tree fall gaps 
and existence of several groups of light-demanding species (e.g., Brazil nut and 
Cecropiaceae). 

We found that the PS forest plots differ structurally from other forests by a 
greater number of stems, the distribution of tree diameter size, and basal area, 
especially the site Limon 3, which has 80 more stems than other plots (Table L). 
In contrast, the TF forest plots in the Palma Real watershed present the typical 
structure in comparison to reports from other regions in Madre de Dios, and 
the Amazon basin, in general (Pitman et al. 2001). 

The PS forests of the Palm Real watershed are characterized by several 
months of anaerobic conditions because of the inundation of the area during 
the rainy season from at least November to February. As suggested by Terborgh 
and Andresen (1998) and Connell and Lowman (1989), who note that swamp 
forests are characterized by low tree diversity and high dominance by one ora 


630 BRIT.ORG/SIDA 22(1) 


low number of species, we expected to find a relatively low species diversity in 
the PS forests of the Palm Real watershed. However, the most significant result 
of this study was that the PS forests in the Palma Real watershed were found to 
have a higher tree species diversity per unit area than TF forests of the region. 
Limon 3,a PS forest plot, was the most diverse of all plots sampled, with 167 tree 
species, only seven species less than the average diversity of 175 species across 
l4 one-hectare TF forest plots sampled in Madre de Dios, Peru, by Pitman et al. 
(2001). Although Oculto 3 was not as diverse as Limon 3, with 126 species it 
had more or just slightly fewer species than the TF forest plots sampled, and it 
was more diverse than the two neighboring TF forest plots from the same site 
(Oculto | and 2). All of our TF forest plots from the Palma Real watershed were 
less diverse than TF forest plots sampled in the neighboring Tambopata region, 
which range between 151 and 197 species (Gentry 1988; Phillips et al. 2003). 

It is interesting to compare the diversity of the two PS forest plots of the 
Palma Real watershed with other one-hectare swamp forest plots in the Madre 
de Dios region. Pitman et al. (1999) and Terborgh and Nufez (unpublished data) 
have found a range of 61-129 tree species among five one-hectare plots in Madre 
de Dios, Peru. Phillips et al. (2003) report 158 tree species from a swamp forest 
plot in the Tambopata region. The Oculto 3 PS forest plot from Palma Real falls 
out at the upper limit of the first range, with 126 species total. Limon 3, with 
167 species, appears to be the most diverse swamp forest plot discovered to date 
in Madre de Dios, Peru. And, as mentioned, Limon 3 is nearly as diverse or is 
more diverse than many non-wetland TF and floodplain forest plots in Madre 
de Dios, Peru. 

Our results and discoveries lead us to some important questions about the 
forests of the Palma Real watershed and argue for continued studies of this re- 
gion. First of all, why are the PS forest plots of Palma Real so high in diversity in 
comparison to plots in neighboring non-swamp forest plots? And second, why 
are the TF forest plots relatively low in diversity compared to the PS forest plots 
of the Palma Real watershed, especially Limon 3, and other TF forest plots from 
the Tambopata region and Madre de Dios, Peru, in general? 

The importance of the PS forest in the region can be seen in the Landsat 
satellite image (Fig. 6), which shows the great expanse of these forests across 
the landscape from the Tambopata River in Peru to the Heath River on the Bo- 
livian-Peruvian border and beyond to the east into Bolivia. Preliminary studies 
in the La Torre River watershed (Cornejo, unpublished data) show a forest with 
the same characteristics of the PS forests of the Palma Real watershed, with a 
dominance in the understory of Lycopodium sp., Selaginella sp., Clusia spp., 
Melastomataceae spp., and other species of non-woody plants that were not 
sampled in our forest plots. Interestingly, such plants are also found in the vast 
wetland savannahs to the east, known as the Pampas del Heath along the bor- 
der of Peruand Bolivia. Local people in the region, in passing conversations with 


peed 


= 


CORNEJO ET AL., FLORISTIC PERU 631 


our team, have mentioned that the Pampas del Heath wetland savannahs can 
develop into dense swamp forests, especially in the absence of management by 
fire. We hypothesize that these diverse PS forests may represent a successional 
stage of vegetation derived from the Pampas del Heath savannahs that are more 
common to the east. This hypothesis could be tested using a combination of 
remote sensing and intensive ground-truthing between the Tambopata region 
and the Pampas del Heath savannahs to the east. The Pampas del Heath sits on 
a thick mass of organic matter that has accumulated over hundreds of years of 
deposition and slow decomposition characteristic of wetland ecosystems (i.e., 
bogs). In other words, perhaps the scattered PS forests of the Palma Real water- 
shed and surrounding region represent a preliminary stage of succession from 
open wetland savannah to tropical forest. Pollen cores could shed some light on 
the succession of these vegetation types. 

This leads us to our second open-ended question, which is why are the TF 
forest plots of the Palma Real relatively low in diversity? The forests of Madre 
de Dios, Peru, are referred to as subtropical moist tropical forests (Foster et al. 
1994). It is possible that the forests in the Palma Real watershed receive less pre- 
cipitation than forests to the west, from Tambopata to Manu, in Madre de Dios, 
Peru. However, there are no climate data available to test this hypothesis. An- 
other possibility is that the soils of TF forests in the Palma Real region are nu- 
trient-poor. We do not have data that enable us to test this hypothesis. We rec- 
ommend future soil sampling in association with vegetation studies in the Palma 
Real region. 

Itis possible that with continued sampling of PS and TF forests in the Palma 
Real watershed, other patterns of plot diversity will be discovered, including 
more diverse TF forests and differences in PS forest diversity. Clearly, more stud- 
ies are needed of PS and TF forests of the Palma Real watershed. Considering 
that the Palma Real watershed is one of the most important zones of Brazil nut 
extraction in all of the southwestern Amazon, we conclude by arguing for more 
sustained and long-term studies of the botanical and ecological diversity, and 
the conservation and management of this region. 


ACKNOWLEDGMENTS 


We would like to thank the Gordon and Betty Moore Foundation and the Ben- 
eficia Foundation for their support of botanical studies in Madre de Dios, Peru. 
We would like to thank Adrian Forsyth and Enrique Ortiz for their support of 
studies of Brazil nut forests in the Palma Real watershed and for their contin- 
ued support of the Brazil Nut Project (Proyecto Castanales) of the Amazon Con- 
servation Association (ACA) and the Asociacion para la Conservacion de la 
Cuenca Amazonica (ACCA), through which this work was originally supported. 
Fieldwork would not have been possible without the logistical support provided 
by ACCA in Peruand specifically the staff of the Brazil Nut Project in Madre de 


632 BRIT.ORG/SIDA 22(1) 


Dios. We thank the board, administration, and staff of the Botanical Research 
Institute of Texas for their constant support of the Andes to Amazon Botany 
Program, through which this manuscript was produced. We thank Sonia Guerra, 
Amanda Neill, Tiana Franklin, and Keri McNew who have supported us di- 
rectly and indirectly in the preparation of this manuscript. We also thank the 
numerous individuals, such as Piher Maceda, Pedro Centeno, Pedro Maceda, 
three Peruvian field assistants who assisted in many aspects of the fieldwork 
required to complete the study. We are grateful to the Ministry of Natural Re- 
sources of Peru (INRENA) for their support of this project and for providing 
permits for research and collections in Madre de Dios, Peru. We also thank 
Asuncion Cano and staff of the San Marcos Natural History Museum, and Car- 
los Reynel and staff of the La Molina University Forestry Department in Lima, 
Peru, for providing herbarium support to the project. 


REFERENCES 


Beck, S.G. 1984. Comunidades vegetales de las sabanas inundazides en el NE de Bolivia. 
Phytocoenologia 12(2/3):321—350. 

Cotwett, R.K. 2005. Estimates: Statistical estimation of species richness and shared species 
from samples. Version 7.5. URL: http://purl.oclc.org/estimates. 

Connett, J.H. and M.D. Lowman. 1989. Low-diversity tropical rain forests: some possible 
mechanisms for their existence. Amer. Naturalist 134:88-119. 

Foster, R., ef AL. 1994. The Tambopata-Candamo Reserved Zone of southeastern Pert: a 
biological assessment. RAP Working Papers, Vol 6. Conservation International. 

Gentry, A.H. 1988. Tree species richness of upper Amazonian forests. Proc, Natl. Acad. Sci. 
85:156-159. 

Gore, NJ.,and R.K. Cowwett. 2001. Quantifying biodiversity: procedures and pitfalls in the 
measurement and comparison of species richness. Ecol. Letters 4:379-391. 

Haase, R. 1990. Plant communities of a savanna in northern Bolivia Il.“palm swamp'’s, dry 
grasslands, and shrubland. Phytocoenologia 18(2/3):343-370. 

KALLIOLA, R., M. PuHakka, J. Sato, H. Tuimisto, and K. RUoKOLAINEN. 1991. The dynamics, distribu- 
tion, and classification of swamp vegetation in Peruvian Amazonia. Ann. Bot. Fennici 
28:225-239, 

Kvist, LP. and G. Neer. 2001. A review of Peruvian flood plain forests: ecosystems, inhabit- 
ants, and resource use. Forest Ecol Managem. 150:3-26. 

Neset, G., L.P. Kvist, J.K. VANCLAY, H. CHRISTENSEN, L. Freitas, and J. Ruiz.2001.Structure and floristic 
composition of flood plain forests in the Peruvian Amazon |. Overstorey. Forest Ecol. 
Managem. 150:27-57. 

Ouiveira, A.D. de and S.A. Mori. 1999. A central Amazonian terra firme forest. |. High tree 
species diversity on poor soils. Biodivers. & Conservation 8:1219-1244. 

Puitups, O.L., R. VAsquez Martinez, P. Nusiez Varcas, A. Lorenzo Monteacuoo, M, CHuspe ZANS, W. 
GALIANO SANCHEZ, A. PENA Cruz, M, TiMANA, M. Yu-Hatta,and S.Rose.2003. Efficient plot-based 
floristic assessment of tropical forests. J. Trop. Ecol. 19:629-645. 


=e 


CORNEJO ET AL., FLORISTIC PERU 633 


Pitman, N.C.A,, J. TersorGH, M.R. Sitman, and P. Nunez V. 1999. Tree species distributions in an 
upper Amazonian forest. Ecology 80:2651-2661. 

Pitman, N.C.A., J.W. TerBorGH, MLR. SiLMAN, P. Nunez V., D.A. Nei, C.E. Ceron, W.A. Paacios, and M. 
AutestiA. 2001. Dominance and distribution of tree species in upper Amazonian “terra 
firme” forests. Ecology 82:2101-2117. 

TerBORGH, J.W. and E. ANnpresen. 1998. The composition of Amazonian forests: patterns at 
local and regional scales. J. Trop. Ecol. 14:645-664. 

Ter Steece, H., D. SABATIER, H. CASTELLANOS, T. VAN ANDEL, J. DUlveNvooRDEN, A.A. DE OLiveirA, R. Ek, R. 
Livwan, P. Maas, and S.A.Moari. 2000. An analysis of the floristic Composition and diversity 
of Amazonian forests including those of the Guiana Shield. J. Trop. Ecol. 16:801-828. 

Ter Steece, H., N.C.A. PITMAN, D. Saparticr, H. CastTeLLAnos, P. Van Der Hout, D.C. Daky, M. SILveiRA, O. 
PHILLIPS, R. Vasquez, T. Van ANDEL, J. DulveNvooRDEN, A.A. DE OLIvEIRA, R. Ek, R. LILWAH, R. THOMAS, J. 
Van Essen, C. BAiDer, P. Maas, S.A. Mori, J. TeR80RGH, P. Nunez Varcas, H.Mocotton, and W. Morawerz. 
2003. A spatial model of tree alpha-diversity and tree density for the Amazon. 
Biodivers.& Conservation 12:2255-277. 

VALENCIA, R., H. Batstev, G. Paz, and C. Mino. 1994. High tree alpha-diversity in Amazonian 
Ecuador. Biodivers. & Conservation 3:21-8. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 


CAMPBELL and LYNN LOUGHMILLER. Updated by Damon E. Waitt. 2006. Texas Wild- 
flowers: A Field Guide (Revised Edition). (ISBN 0-292-71286-3, pbk.). University 
of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: http:// 
www.utexas.edu/utpress, 512-471-4032). $19.95, 278 pp., 381 color photo, 
ay Tee, 

This is the successor to the original from 1984, which, 

top three best sellers for the University of Texas Press, with more tha 


edition includes an additional 61 species (for a total of 370), but most rai photos ae been re- 
1, consistently 


ae ere eae 
notes g the 


1155, favayay oy 14 he new 


see Ongne text is mostly pncneneee: The family order i g I 
iff | lated for species names, about 


sing © -aceae’ ‘endings for the 
: percent of which have been supplanted or modified since 1984. Del Damon Waitt of the Wild- 
flower Center in Austin has been responsible for the taxonomic revision and accuracy of the photo 
identifications, as both of the Loughmillers have passed on 
In explanation of why faites 2 I sé 1 lly appearance from the first,” the intro- 
duction notes that the cover stock is durable and the en narrower—changes that are “part of 
the University of Texas Press plan to develop a complete series of natural history field guides.” Ap- 
parently as part of the same revamp, margins are pee spaces expanded oa the words and 
| less COMIPAacl COX. The new | SAaY¥e consid- 


lines, and the font less bold—resulting in 
erably smaller and seem subsidiary to the text. Edition 2 is useful, but if you don’t have the first, try 
to find a copy of it also, since the original Loughmiller photos, in addition to their larger size, often 


here it was 


show views more representative of the species, and a brief statement for V 
taken (e.g., “Photographed near Blanco State Park in April,” “Photographed in the Big Thicket in 
May”).—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, 
USA 


SIDA 22(1): 634. 2006 


A VASCULAR PLANT INVENTORY OF STARKEY 
WILDERNESS PRESERVE, PASCO COUNTY, FLORIDA 


Emily Ferguson Richard P Wunderlin 
2531 River Plaza Dr, #52 Department of Biology 
Sacramento, California 95833, U.S.A. University of South Florida 


Tampa, Florida 33620-5200, U.S.A. 


ABSTRACT 


Starkey Wilderness Preserve d and 1 by the Southwest Florida Water Management Dis- 
trict, contains 7,739.5 hectares with 18 slanee communities. A floristic inventory was conducted on 
449.2 hectares within the property which consists of 11 community types and is considered as rep- 
resentative of the entire Preserve. The objective of this study, conducted from May 2003 to October 
2004, was to compile a list of the vascular plant taxa found to be used by the Southwest Florida 
ter Management District to help in their management regimes. A total of 483 taxa were collected, 
representing 104 families, and 271 genera. Of these, 444 are native taxa, 16 Florida endemic taxa, 39 
non-native taxa, 12 encaneciess threatened or commercially exploited taxa, and 33 county records. 
Each is described and an annotated list of the vascular plant taxa is presented 


pi 


RESUMEN 


La reserva de Starkey Wilderness, gestionada por el Southwest Florida Water Management District, 
tiene 7,739.5 hectareas con 18 aeaaae vegetales. Se realiz6 un inventario floristico en 449.2 
hectareas de la reserv 1 tipos de comunidades y que se consideran representativos de 
la Reserva entera. El cbeiv0 us este ae realizado desde mayo de 2003 hasta octubre de 2004, 


fue compilar una lista d I sadas por el souces Florida Water Manage- 
ment District para ayuda égi | ion. Se col de 483 taxa, que ce 
104 familias, y 271 géneros. De slice 444 ¢ son taxa nativos, 16 taxa endémicos de Florida, 39 taxa 


aldctonos, 12 taxa en peligro, amenazados o explotados comercialmente, y 33 citas del ois Se 
describen las comunidades vegetales y se presenta una lista comentada de los taxa de plantas 


vasculares 
INTRODUCTION 


Florida is a unique state with a relatively flat topography and generally fast 
draining soils. These conditions would presumably lead toa flora with low di- 
versity, but that is not the case. The state is about 1000 km long and spans about 
6.5 latitude resulting in a diverse flora consisting of nearly 4,200 taxa of native 
and naturalized non-native vascular species (Wunderlin & Hansen 2003), the 
third most diverse in the United States. As Florida is also the fourth most popu- 
lated state, it is important to work to conserve and protect the state’s flora. 
Several agencies in Florida work to secure large tracts of landinan effort to 
prevent development and to establish areas of conservation. One of these, the 
Southwest Florida Water Management District (6WFWMD), owns many par- 
cels of land for a variety of water management practices including the protection 


SIDA 22(1): 635 — 659. 2006 


636 BRIT.ORG/SIDA 22(1) 


and development of potable water supplies, aquifer recharge, water quality en- 
hancement, restoration and protection of natural systems, and structural flood 
control. The property in which the inventory was conducted is owned and 
managed by SWFWMD. The results of this research are intended to help 
SWEWMD make knowledge-based management plans for its property. 


Site Overview 

Physical Location.—Starkey Wilderness Preserve is located in southwestern 
Pasco County, Florida, near the west coast of central Florida (Fig. 1). It is 6.4 km 
northeast of Seven Springs and 4.4 km northwest of Odessa. Main access to the 
Preserve is Wilderness Road approximately 5.6 km from State Road 54. A por- 
tion considered to represent the major habitats found within the entire Preserve 
was selected for this inventory. The Starkey Wilderness Preserve is included 
within Sections 1-17, 21-24, and 26-27, Township 26 South, Range 17 East and 
Sections 10-11, 13-15, 22-27, and 33-36, Township 25 South, Range 17 East. The 
entire Preserve comprises approximately 7,739.5 hectares. onsists of 18 plant 
communities. Of these, 11 occur within the study site. The study site, consisting 
of 449.2 hectares, includes portions of Sections 3-4, 8-10, and 15-17, Township 
26 South, Range 17 East. The boundaries of the study site comprise a power line 
artery along the western edge, a paved biking trail at the north, an unpaved 
road running due south along the eastern edge, and the Anclote River along the 
south. 

Early Inhabitants.—The first humans who lived near the Starkey Wilder- 
ness Preserve were the Safety Harbor Culture or the Tocobaga, who inhabited 
the Florida Gulf coast (Lawson et al. 1981). The period of their inhabitance was 
from 900 A.D. through the early eighteenth century (Milanich 1994). The settle- 
ment area extended from the Withlacoochee River (north and east boundary 
of Citrus County) south to the Charlotte Harbor area (Charlotte County). 
Tocobaga arrowheads have been unearthed on the Starkey property, suggest- 
ing their use of the land to hunt (Lawson et al. 1981). Approximately 33 prehis- 
toric archaeological sites have been identified as temporary campsites for hunt- 
ing within the Starkey Wilderness Preserve (GSWFWMD 1990). Hunters on the 
Preserve could have originated from two different sites. The first, a flint work- 
shop located 3.2 km north of the Anclote River along the coast; and the second, 
Tarpon Springs at the mouth of the Anclote River (Willey 1949). In the eigh- 
teenth century, after the Spanish established themselves in much of the state, 
the remaining Tocobaga are believed to have joined the Creek tribes from fur- 
ther north as they moved south seeking refuge from persecution by the new 
settlers. 

The land now included in the Starkey Wilderness Preserve passed from 
the Federal Government to the State of Florida and private ownership beginning 
in 1856 (Lawson et al. 1981). Principal land uses were for turpentine production 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA 


Soil types oN 
QB Adamsville fine sand C 
— Basinger fine sand 

| Cassia fine sand 
— Chobee soils 

[3] Immokalee fine sand 
___] Myakka fine sand 
2 Paola fine sand 
Pomello fine sand 
MM Pomona fine sand 
fT Sellers mucky loamy soil f 
| Smyrna fine sand : 
(99 Wauchula fine sand 


0 250 = 500 1,000 1,500 2,000 
oo = J Mer Meters 
Fic. 1. A. Location of the Jay B. Starkey Wild P , Pasco County, Florida. B. Jay B. Starkey Wilderness Preserve 
ith +h f+h tod C pl re ary 


Jay B. Starkey Wilderness Preserve study area. 


638 BRIT.ORG/SIDA 22(1) 


and for cypress and pine lumber. In 1937, Jay B. Starkey and his partners, Ernest, 
Dave, and Howard Cunningham, bought 6,070 hectares. The C-S (Cunningham- 
Starkey) Ranch was used to raise cattle by both families. As land values in- 
creased and cattle yields decreased, portions of the land were sometimes sold 
to developers. Jay B. Starkey, who outlived his partners, finally sold the cattle 
and the equipment to his son Jay. B. Starkey, Jr. The senior Starkey still lived and 
worked the land with his son for cattle ranching, and later during World War 
IL. converted about 30 hectares into an orange grove. Over the years, items such 
as timber, pine stumps, and resin were sold to provide extra income. In 1975, as 
the population increased in the county, Jay B. Starkey, Sr. decided to preserve 
the land from development by selling part of the ranch to SWFWMD with the 
understanding that SWFWMD would maintain the land in its natural state 
(Lawson et al. 1981). By 1980, SWFWMD had acquired 1,407.9 hectares (Wells 
2004). The SWEWMD has purchased additional property since then and the 
Starkey Wilderness Preserve now comprises approximately 7,739.5 hectares. 

Climate.—The climate of Pasco County is humid and subtropical (Chen & 
Gerber 1990). The constant cyclical pattern consists of cool, dry winters and 
warm, humid summers. Annual mean temperature is 22°C (SWF WMD 1988). 
During the winter months, December through February, the monthly mean 
temperature is 16°C. The temperature will usually drop to freezing at least once 
a year. During the summer months, from June to September, the monthly mean 
temperature is about 27° C, with highs reaching about 33°C. 

Rainfall in Pasco County varies seasonally, oscillating between the rainy 
season and the dry season (SWF WMD 1988). From October to May, rainfall is 
less than 10.2 cm per month, while during the summer months, June through 
September, the area receives about 18.4 cm of rainfall per month. This accounts 
for about half of the rainfall for the year. In spring, central Florida, including 
Pasco County, experiences harsher and longer, dry drought-like conditions than 
either northern or southern Florida (Chen & Gerber 1990). The high amounts 
of rain that fall from June to September are attributed to the convective clouds, 
sea breezes, and tropical storms. By the beginning of October, the rainfall de- 
creases by about 50 percent from that of the summer months, indicating the 
beginning of the dry period. 

Geology.—The geology underlying the Starkey Wilderness Preserve is the 
Suwannee Formation deposited during the Oligocene epoch of the Tertiary 
period (34 to 24 MYBP) (Scott et al. 2001). This formation is a yellow to white, 
fossiliferous, fine-grained limestone (Wetterhall 1964). It is a very porous for- 
mation that contains the Upper Floridan Aquifer. Below the Suwannee Forma- 
tion are the Ocala and Avon Park Formations, both deposited during the Eocene 
epoch of the Tertiary period (55 to 34 MYBP). Both of the lower formations are 
also important parts of the Floridan aquifer. 

Soils.—Twelve distinct soil types occur within the study site. The domi- 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA 639 


nant types are: Chobee, frequently flooded (22%), Adamsville fine sand (19%); 
Myakka fine sand (13%); Immokalee fine sand (11.5%); and Sellers mucky loamy 
fine sand (10%). Other soil types present are Pomona fine sand (8%); Pomello 
fine sand, 0 to 5 percent slope (6%); Wauchula fine sand, 0 to 5 percent slope 
(4%); Cassia fine sand, 0 to 5 percent slope (3%); Smyrna fine sand (1.5%); Paola 
fine sand (1%); and Basinger fine sand (1%) (Stankey 1982). 

Topography and Hydrology.—Starkey Wilderness Preserve is located in the 
Gulf Coastal Lowlands physiographic province, which is characterized by low 
elevations and poor drainage. Within the study site, the elevation ranges from 
9.1 to 16.8 m (USGS 1974). The majority of the site has an elevation of 10.7 to 12.2 
m. The swamp and river floodplain area ranges from 9.1 to 10.7 m. 

The Anclote River, located primarily in Pasco County, is 27.4 km long. 
About 8.1 km of the Anclote, excluding the south branch, is within the Starkey 
Wilderness Preserve (SWFWMD 1990). The southern border for the study site 
is an approximately 3.2 km section of the Anclote River (fig. Ll). The Anclote is 
characterized as a slow moving, turbid, acidic, black water stream. The chan- 
nel is not more than 6 m wide at any point along the southern boundary of the 
study site. During low flow stage, the stream is confined to the channel. As the 
rainy season starts, the Anclote overflows its banks inundating the floodplain 
forest, and remains at flood stage for two to three months. When the river over- 
flows its banks, the water flowing into the floodplain removes waste and pro- 
vides nutrients to the floodplain community. The sediment load settles out as 
the water moves out of the channel. Heavier sediments are deposited first closer 
to the channel due to their weight, and finer, lighter sediments settle out more 
slowly further away from the channel. 

Disturbances Affecting the Study Site—With the exception of the effects 
of cattle grazing, lumbering, and other minor events, the study area has under- 
gone few changes. The western boundary of the study site is the right-of-way 
for the Florida Power Corporation electrical transmission line. This 46 m wide 
artery was constructed in the 1970s (SWFWMD 1990). In order to maintain the 
right-of-way the area is frequently mechanically cleared. In addition, SWF WMD 
built 14 active production water wells within the Preserve. These wells pump 
water from the Preserve to provide potable water for the city of New Port Richey. 
Three production wells occur within the study site. An approximately 3,240- 
hectace portion of the Preserve, the Jay B. Starkey Wilderness Park, is open to 
the public for a number of recreational opportunities. In order to provide these 
recreational activities, SWFWMD constructed a paved road, hiking/biking 
trails, horse riding trails, primitive camp sites, and semi-primitive road beds. A 
number of these features occur crinee the study site as well. 

The most recent disruption within the study site was a wildfire that was 
believed to have started from a lightning strike June 3, 2004. After persisting 
through the night, the wildfire consumed approximately 162 hectares on June 


— 


640 BRIT.ORG/SIDA 22(1) 


4, primarily in a band through the center of the study site. This fire occurred 
towards the end of the research collection period and thus had little impact on 
data collection. 

Management Regimes.— The management plan devised and implemented 
by SWF WMD (1990) includes several conceptual land uses. The most impor- 
tant function for the District is to manage the well field withdrawals. The con- 
sumptive use permit renewed in 1979 allows the city of New Port Richey to 
withdraw 30 to 57 million liters per day from the 14 wells within the Preserve 
(SWFWMD, 1990). This water withdrawal has produced shorter hydroperiods, 
lower peak water levels, increased rates of succession into wetland areas, and 
unusual plant associations (SWFWMD 1990). 

To manage areas of historical significance, SWF WMD protects them from 
disturbance by keeping hiking and horse-riding trails away from them. 
SWFWMD has opened the Preserve for “low intensity resource-based” recre- 
ation by building facilities for local use (SWFWMD 1990). These recreational 
facilities are managed by Pasco County. They include hiking, biking, horse 
riding, birding, camping, and any other non-motorized activities. The hiking/ 
biking trails are separate from the horse riding trails to avoid safety hazards. 

Land management practices exercised by SWFWMD include a number of 
ways to improve or preserve the quality of the property. The District uses pre- 
scribed burning to conserve various communities and to restore habitats. Dur- 
ing the summer months, when there is greater thunderstorm activity, higher 
lightning frequency giving way to natural wildfires which SWFWMD contains 
and controls. The District normally burns during the winter months in order 
to limit and control fires (SWFWMD 1990). 

A population of feral hogs has disturbed the soil while rooting, causing 
damage to some habitats and previously preserved historical sites. Part of the 
management plan is to reduce or minimize their impact. 


METHODS 


Vascular plant specimen vouchers were collected from May 2003 to October 
2004 using standard field collection and herbarium techniques. Notes were 
made for each collection regarding the habitat and frequency of occurrence. 
One complete set of voucher specimens was deposited at the University of South 
Florida Herbarium (USF). The floras of Wunderlin (1998) and Wunderlin and 
Hansen (2003) were utilized for identifications, followed by verification with 
specimens in the USF herbarium. 

Vascular plants of special interest include those taxa that are Federal and/ 
or State listed species and exotic species listed by the Florida Exotic Pest Plant 
Council (FLEPPC 2005). Species unreported for Pasco County as determined by 
Wunderlin and Hansen (2005) are noted. 

The plant community classification for the study site follows that of Florida 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA 641 


Natural Areas Inventory (FNAI1990). The plant communities are described and 
include the dominant species found within each layer (overstory, understory, 
and herbaceous ground layer). 


RESULTS AND DISCUSSION 


A total of 483 taxa were collected within the study site within Starkey Wilder- 
ness Preserve. The flora is comprised of 104 families and 271 genera (Table D). 
The families with the greatest number of taxa are: Ast (66), Poaceae (62), 
Cyperaceae (39), and Fabaceae (28). The genera with the most abundant taxa 
are: Rhynchospora (12), Quercus (10), Hypericum (9), Andropogon (8), Ludwigia 
(8), and Dichanthelium (8). 

About 92% of the flora in the study site is native. Sixteen (3%) of the spe- 
cies found are endemic to Florida. Four of the 39 non-native taxa are listed by 
the Florida Exotic Pest Plant Council (FLEPPC 2005). Thirty-three taxa are 
records for Pasco County (Wunderlin & Hansen 2005). Twelve taxa are listed as 
endangered, threatened, or commercially exploited in Florida (Coile & Gar- 
land 2003). No Federally listed taxa occur on the site. 

Starkey Wilderness Preserve consists of 18 plant communities. Eleven of 
these occur within the study site. Comparing the Preserve with the study site 
shows the study site is representative of the Preserve (Table 2). The plant com- 
munities that fall outside the boundaries of the study site are not dominant ones 
and contribute only 4% of the total area of the Preserve. Thus the plant list com- 
piled for the study site is fairly representative for the majority of the Preserve. 


NATURAL COMMUNITIES 


The classification for plant communities found within the study site following 
the Florida Natural Areas Inventory guidelines (FNAI 1990, 2004) are discussed 
below from the lowest elevation type to the highest followed by the ruderal areas. 

Floodplain Forest.—The floodplain forest is a forested wetland that con- 
sists of 64.3 hectares adjacent to the River. Flooding is closely linked to the rain 
events that occur throughout the summer months. The soil is Chobee (Stankey 
1982) 

The dominant tree species that make up the overstory of the floodplain 
forest are Acer rubrum, Carpinus caroliniana, Fraxinus caroliniana, Ilex cassine, 
Nyssa sylvatica var. biflora, Persea palustris, Quercus laurifolia, Quercus 
virginiana, Sabal palmetto, Taxodium distichum, Ulmus alata, and Ulmus 
americana. Some of these are buttressed due to the long hydroperiods. Note- 
worthy epiphytic species are Encyclia tampensis, Epidendrum conopseum, 
Pleopeltis polypodioides, and Tillandsia spp. The woody understory includes 
Berchemia scandens, Campsis radicans, Cephalanthus occidentalis, Psychotria 
nervosa, Psychotria sulzneri, and Toxicodendron radicans. The herbaceous 
ground layer consists of a number of pteridophytes including Blechnum 


642 BRIT.ORG/SIDA 22(1) 


serrulatum, Osmunda cinnamomea, Osmunda regalis, Thelypteris interrupta, 
Taste 1. Synopsis of the vascular plants found in the Jay B. Starkey Wilderness Preserve study area. 


Taxa Genera Families Endemics Non-native Co. Records 
Pteridophytes 13 9 7 0 1 2 
Gymnosperms 7 4 3 0 0 0) 
Monocotyledons 164 71 24 2 17 14 
Dicotyledons 299 187 70 14 21 17 
Totals 483 27) 104 16 39 33 


Taste 2, Summary of the plant communities in the Jay B. Starkey Wilderness Preserve and the study 
area. 


Plant Community SWP SA Plant Community SWP SA 
Basin Marsh 101.2 Oak Scrub 70.4 

Basin Swamp 1,422.1 61.5 Pine Plantation 22.2 

Clastic Upland Lake 10.9 Ruderal and Disturbed = 888.3 0.4 
Depression Marsh 48.6 2.0 i Pine Scrub 206.0 20.2 
Dome Swamp 2347 10.5 dhill 275.6 344 
Dry Prairie 0.8 ate Flatwoods 3314 40.5 
paieaiee Forest 746.2 64.3 Wet Flatwoods 79.3 

Hydric Hammock 28.3 Wet Prairie 40.9 41 
Mesic Seen 3,005.6 207.2 Xeric Hammock 227.0 4.1 


Wood wardia areolata,and Woodwardia virginica. aa common are Carex gigan- 
tea, Hypoxiscurtissii, [ris hexagona, Rhyncl 


i Saururuscernuus, Scleria 
triglomerata, and Spiranthes odorata. A non-native species found along the 
unimproved trails into the floodplain forest is Urena lobata. 

Basin Swamp.—A basin swamp is a large irregularly-shaped depression 
that is not part of a river, although during the rainy season it may be connected 
via surface flow to the river. This region is vegetated by species similar to the 
floodplain forest. The soils in the study site are Chobee and Sellers mucky loamy 
fine sand (Stankey 1982); both of these are frequently inundated with water. 
The study site contains 61.5 hectares of basin swamp. The largest of this com- 
munity type in the study site is located in the middle of the lower half of the 
site; it was probably an oxbow originating from the Anclote River. 

The dominant overstory species include Acer rubrum, Carpinus 
caroliniana, Diospyros virginiana, Fraxinus caroliniana, Gordonia lasianthus, 
Ilex cassine, Nyssa sylvatica var. biflora, Persea palustris, laxodium distichum, 
Ulmus alata, and Ulmus americana. The dominant woody understory species 
are Cephalanthus occidentalis, Lyonia lucida, and Myrica cerifera. The herba- 
ceous ground layer consists primarily of Amphicarpum muhlenbergianum, 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 643 


Carex spp., Cladium jamaicense, Cyperus spp., Eleocharis spp., Eriocaulon 
decangulare, Juncus spp., Leersia hexandra, Panicum hemitomon, Peltandra 
virginica, Pontederia cordata, Rhexia spp. Sagittaria lancifolia, Saururus 
cernuus,and Scleria spp. The basin swamp also contains a number of epiphytic 
species such as Pleopeltis polypodioides, Tillandsia recurvata, Tillandsia 
simulata, Tillandsia usneoides, and Tillandsia xfloridana. 

Dome Swamp.—Dome Swamps are found throughout the study site. The 
study site contains 10.5 hectares of dome swamp. These circular to oblong de- 
pressions contain Sellers mucky loamy fine sand, a very poorly-drained soil 
type (Stankey 1982). An impermeable clay layer beneath the soil helps main- 
tain water levels in these depressions. The depression is due to the soils slump- 
ing into sinkholes formed in the limestone rock. Most of the water found in the 
swamp is surface runoff from surrounding upland communities. The dome 
swamp may become completely desiccated at the end of the dry season, expos- 
ing the peat layer for a couple of weeks. These areas have a domed profile with 
smaller trees in the shallow water near the edge and larger trees in the deeper 
water in the middle of the swamp. 

The dominant tree in the dome swamp is Taxodium ascendens. Other 
prominent overstory species include Acer rubrum, Diospyros virginiana, Ilex 
cassine, Nyssa sylvatica var. biflora,and Persea palustris. Typical woody under- 
story species are Lyonia lucida, Myrica cerifera, Smilax laurifolia,and Toxico- 
dendron radicans. The herbaceous ground layer includes the pteridophytes 
Thelypteris interrupta, Woodwardia areolata,and Wood wardia virginica. Other 
ground layer species are Drosera capillaris, Helenium pinnatifidum, Polygala 
cymosa, Polygonum hydropiperoides, Rhynchospora latifolia, Spiranthes 
laciniata,and Xyrisspp. The epiphytic flora is similar to that of the floodplain 
forest and basin swamp and includes Pleopeltis polypodioides, Tillandsia 
recurvata, lillandsia simulata, Tillandsia usneoides, and Tillandsia xfloridana. 
A rare find in one of the swamps was Vittaria lineata. 

Depression Marsh.—Depression marshes are characterized by low depres- 
sions in the flatwoods ity due to slumping around the edges of a sink- 
hole or water collecting on top of a hardpan subsurface layer. Two hectares of 
depression marsh occur in the study site. The soils are Adamsville fine sand 
and Basinger fine sand (Stankey 1982). 

These regions typically do not contain any overstory species. Woody spe- 
cies include Hypericum fasciculatum, Ilex gla bra, Myrica cerifera, and Stillingia 
aquatica. The herbaceous ground layer contain hicarpum muhlenbergianum, 
Carex spp., Cyperus spp., Eleocharis spp., Eriocaulon decangulare, Juncus spp., 
Lachnanthes caroliana, Panicum hemitomon, Rhexia spp., Sabatia grandiflora, 
Scleria spp.,and Xyris spp. 

Wet Prairie.—About 4.1 hectares of the study site is covered by this open 
wetland herbaceous community. It is often found within the mesic flatwoods 


644 BRIT.ORG/SIDA 22(1) 


or surrounding the fringe of a dome or basin swamp. The soils are Basinger fine 
sand and Myakka fine sand (Stankey 1982). The area is poorly drained due to 
the subsoil layer of hardpan clay. 

The wet prairie includes some native invading woody species: Hypericum 
fasciculatum, Hex glabra, Myrica cerifera,and Stillingia aquatica. The herbaceous 
ground layer primarily consists of Aletris lutea, Amphicarpum muhlenbergianum, 
Aristida stricta, Carex spp., Cyperus spp., Drosera capillaris, Eleocharis spp., 
Eriocaulon decangulare, Eupatorium mohrii, Fimbrystylis spp., Juncus spp., 
Lachnanthes caroliana, Panicum hemitomon, Polygala spp., Rhexia mariana, 
Sabatia spp., Scleria spp., Syngonanthus flavidulus, and Xyris spp 

Mesic Flatwoods.—T he most dominant fire adapted community found in 
the study site (207.2 hectares) is mesic flatwoods. This community has a rela- 
tively flat topography. The dominant soils are Immokalee fine sand, Myakka fine 
sand, and Smyrna fine sand (Stankey 1982). These soils normally have low levels 
of nutrients and organic matter. A hardpan clay layer under the soil causes poor 
water drainage in the rainy season and xeric conditions during the dry season. 

The overstory layer is composed of Pinus elliotti and Pinus palustris. The 
pines are widely spaced across the community allowing for a dense woody un- 
derstory layer. The understory layer includes Ilex glabra, Lyonia lucida, and Ser- 
enoa repens. Smaller understory layer species are Gaylussacia frondosa, Quercus 
minima, Vaccinium corymbosum, and Vaccinium myrsinites. The rich herba- 
ceous ground layer includes Amphicarpum muhlenbergianum, Aristida stricta, 
Dichanthelium ensifolium, Dichanthelium portoricense, Elephantopus elatus, 
Eupatorium mohrii, Euthamia caroliniana, Pityopsis graminifolia, Polygala 
lutea, Polygala setacea, Pteridium aquilinum, Pterocaulon pycnostachyum, 
Seymerid cassioides, and Sorghastrum secundum. 

Sandhill.—The sandhill community makes up 34.4 hectares of the study 
ite. There are two areas; both are relatively high in elevation at 12.2-15.2 m and 
have topography consisting of rolling hills and gentle slopes. The soil of both 
areas is Adamsville fine sand (Stankey 1982), which is also found under the 
scrubby flatwoods community within the study site. The deep sandy soils that 
are easily leached and well drained create xeric characteristics in this commu- 
nity. The overstory layer is sparse, allowing sunlight to reach the ground layer, 
adding to the xeric condition. 

The dominant overstory species include Pinus palustris, Quercus incana, 
Quercus laevis, and Quercus margaretta. Understory species include Asimina 
obovata, Asimina reticulata, Diospyros virginiana, Gaylussacia dumosa, Licania 
michduxii, Myrica cerifera, oes minima, Rhus Se and Serenod 
repens. The herbaceous ground cover layet consists pl imat ily of Aristida stricta, 
Asclepias humistrata, Balduina angustyold. Dichanthelium portoricense, 
Phoebanthus grandiflorus, Pityopsis graminifolia, Pterocaulon pycnostachyum, 
Rhynchospora megalocarpa, and Sorghastrum secundum. 


— 


WY 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 645 


Scrubby Flatwoods.—This community consists of 40.5 hectares within the 
study area. The soil is Adamsville fine sand (Stankey 1982). The vegetation con- 
sists of a mixture of mesic flatwoods and sand pine scrub species. It is charac- 
terized by an open pine canopy with sparse scattered clumps of oak species 
and patches of open sand. 

The species found in the overstory include Pinus clausa, Pinus palustris, 
Quercus chapmanii, Quercus geminata, and Quercus myrtifolia. The dense un- 
derstory layer contains Garberia heterophylla, Lyonia ferruginea, Lyonia 
fruticosa, Serenoa repens,and Ximenia americana. The herbaceous ground layer 
contains Aristida spiciformis, Aristida stricta, Carphephorus corymbosus, Dichan- 
thelium portoricense, Lupinus diffusus, Pteridium aquilinum,and Rhynchospora 
megalocarpa. Several Pinus palustris have the epiphyte Tillandsia xfloridana. 

Sand Pine Scrub.—One area containing a scrub community totaling 20.2 
hectares occurs within the study site along the western boundary. This com- 
munity develops along the ridgeline of ancient dune formations composed of 
very fine sand sometimes referred to as “sugar sand.” The soil is Pomello fine 
sand (Stankey 1982). These fine sands allow rain water to quickly percolate 
down to the aquifer creating a xeric growing condition. The scrub can have an 
open or closed canopy consisting of Pinus clausa with various oaks and shrubs 
dominating the understory. Normally this community type exhibits patches 
of exposed sand; however, in the study site these are very few. The scrub found 
within the study site has mature Pinus clausa, an indication of fire suppres- 
sion. 

Under the Pinus clausa, the dominant overstory also contains Quercus 
chapmanii, Quercus geminata, and Quercus myrtifolia. The understory shrub 
layer contains Asimina obovata, Ceratiola ericoides, Garberia heterophylla, 
Licania michauxii, Lyonia ferruginea, Lyonia fruticosa, Serenoa repens, and 
Ximenia americana. The herbaceous ground cover is sparse; however some com- 
mon herbs found there are Palafoxia intergrifolia, Pityopsis graminifolia, and 
Rhynchospora megalocarpa. 

Xeric Hammock.— Xeric hammock is generally considered a degraded fire- 
excluded scrub or sandhill community. Thi unity in the study site is an 
advanced sandhill community and consists of 4.1 hectares on the fringes of the 
northern sandhill community. The xeric hammock occurs on two soil types, 
Adamsville fine sand and Immokalee fine sand (Stankey 1982). The commu- 
nity lacks a closed overstory canopy, but a low canopy is present from the shrub 
layer dominated by oaks 

The overstory species consist of Pinus palustris (few), Quercus geminata, 
and Quercus laevis. The dense woody understory consists primarily of Licania 
michauxii, Lyonia fruticosa, Quercus chapmanii, Quercus myrtifolia,and Serenoa 
repens. The herbaceous ground es Sue ae stricta, Dichanthelium por- 
toricense, Galactia elliottii, Pteri mand Rhynchospora megalocarpa. 


L 


646 BRIT.ORG/SIDA 22(1) 


Ruderal.—Ruderal areas within the study site include regions of distur- 
bance due to anthropogenic changes. This encompasses 0.4 hectare of road- 
sides, hiking trails, horse riding trails, well pump houses, improved pasture, and 
the fringe of the power line artery. These areas contain a high number of non- 
native species because of the recent disturbances caused by their construction 
and subsequent use. 

The most prominent non-native is Paspalum notatum followed by 
Crotalaria spp., Indigofera hirsuta, and Ludwigia peruviana. The Southwest 
Florida Water Management District (SWFWMD) has worked to manage these 
ruderal areas to encourage recruitment of native species. 

ANNOTATED LIST OF THE VASCULAR FLORA 
The vascular flora of the study site within the Starkey Wilderness Preserve in 
the following list is documented by vouchered specimens in the University of 
South Florida herbarium (USF). The list is divided into four major sections: ptert- 
dophytes, gymnosperms, monocots, and dicots (basal angiosperms and 
eudicots). Within these sections the list is arranged alphabetically by family, 


genus, and species. The nomenclature follows Wunderlin and Hansen (2003, 
2005). 

Each species and infraspecific taxon is followed by a common name and 
its frequency of oes The frequency of occurrence is given as: R, rare 
less than four individuals found; O, occasional, between four and 15 plants aoe 
and C, common, more than 15 plants in the immediate area. Following the fre- 
quency of occurrence, the primary plant community from which the . 
was collected is noted. The plant communities include: basin swamp (BS); de- 
pression marsh (DM); dome swamp (DW); floodplain forest (FF); mesic flatwoods 
(MF); ruderal (RD); sandhill (SH); scrubby nee (SF); sand pine scrub (SP); 
wet prairie (WP); and xeric hammock (XH). Following this, taxa listed as en- 
dangered, threatened, or commercially exploited in Florida (Coile & Garland 
2003) are noted. The number(s) in brackets at the end of each species are the 
collection number(s) of the first author. 

Species endemic to Florida are listed in bold font. Non-native species are 
marked with an asterisk (*). New records for Pasco County are indicated by an 
underline. Exotic species listed by the Florida Exotic Pest Plant Council (FLEEPC 
2005) are listed with their ranking: Category I—species altering Florida’s natu- 
ral plant community by displacing native species and changing the structure 
of the community; Category I[—species having the ability in the future to alter 
Florida’s natural plant communities by displacing natives and changing the 
community structure. 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 647 


PTERIDOPHYTES 


Blechnaceae 

Blechnum serrulatum Rich.—toothed midsorus 
fern; C; DS (116, 498, 538) 

Woodwardia areolata (L.) T. Moore—netted 
chain fern; C; FF (552, 822) 

Woodwardia virginica (L.) Sm.—Virginia chain 
fern; C; FF (523, 499, 418) 


SL haa a 

Pteri aquilinum (L.) Kuhn © var. 
ae eee (Clute) Clute ex A. Heller— 
tailed bracken fern; C; MF (249, 823 

Lycopodiaceae 

Lycopodiella alopecuroides (L.) Cranfill—foxtail 
club-moss; R; BS (266) 


Osmundaceae 
Osmunda cinnamomea L.—cinnamon fern; C;FF; 
commercially exploited (347) 
Osmunda regalis L. var. spectabilis (Willd.) A 
ray—royal fern; O; FF; commercially ex- 
ploited (553,655) 


— 


Polypodiaceae 

Poe aureum (L.) J. Sm.—golden poly- 
pody; O; DS (579) 

Pleopeltis pobpodotes (L.) E.G. Andrews & 
Windham var. michauxiana (Weath.) E.G. 
Andrews & on eee fern; C; 
FF (398) 


Thelypteridaceae 

*Thelypteris dentata (Forssk.) E.P. St. Jonn— 
downy maiden fern; O; FF (803) 

lyoteris interrupta (Willd.) K.lwats.—hottentot 
fern; C; FF (354) 

Thelypteris kunthii (Desv.) C.V. Morton—wide- 
spread maiden fern; C; FF (554) 


The 


— 


Vittariaceae 
Vittaria lineata (L.) Sm.—shoestring fern; R; DS 
(765) 


GYMNOSPERMS 
Cupressaceae 
Juniperus virginiana —red cedar; R; RD (782) 
Taxodium lens Brongn.—pond-cypress;C; 


S (170) 
Taxodium distichum (L.) Rich.—bald-cypress; C; 


Pinaceae 
Pinus clausa (Chapm. ex Engelm.) Vasey ex 
rg.—sand pine; C; SF (290) 

Pinus elliottii Engelm.—slash pine; C; MF (595) 

Pinus palustris Mill—longleaf pine; C; MF (581) 

Zamiaceae 

Zamia pumila L.—Florida arrowroot;R; MF;com- 
mercially exploited 


— 
Ww 
— 
oO 

= 


MONOCOTS 


Agavaceae 
Yucca filamentosa L.—Adam’s needle; O; SH (91) 


Alismataceae 

Sagittaria graminea Michx. var. graminea— 

15, 380) 

ee a a inea Michx. var. chapmanil J.G. 
d; O; FF (480) 


grassy arro\ shead:O; DS (1 


rrowhea 


—Chapman’'s a 


a ittaria lameifal ia L—bulltongue arrowhead; 


Amaryllidaceae 

Zephyranthes atamasca (L.) Herb. var. treatiae (S. 
Watson) Meerow—Treat’s rainlily; R; FF; 
renee (665) 


Araceae 

Arisaema Hak (L.) Schott—Jack-in-the- 
pulpit; R; FF (740 

Lemnava liane, Phil—Valdivia duckweed; C;BS 
2\5 


Peltandra virginica (L.) Schott—green arrow 

arum; R; BS (776) 
ecaceae 

Sabal minor (Jacq.) Pers.—dwarf palmetto; C; FF 
316 

Sabal palmetto (Walter) Lodd. ex Schult. & 
Schult.£.—cabbage palm; O; FF (794) 

Serenoa repens (W. Bartram) Small—saw pal- 
metto; C; MF (80) 


Bromeliaceae 
Ts if y (ret L 


494) 
Tillandsia recurvata (L.) L—ballmoss; C; SH (270) 
Tillandsia setacea Sw.—southern neecleleaf; C; 

FF (751) 
Tillandsia simulata Small—airplant; C; FF (305, 


= 
— 


Tillandsia usneoides (L.) L—Spanish moss; CG; SH 


(452) 


648 


Tillandsia utriculata L.—giant airplant;O; SH; en- 
dangered (837) 
Tillandsia Xfloridana (L.B. Sm.) H. Luther; C; SF 


Burmanniaceae 
Burmannia capitata (J.F.Gmel.) Mart.—southern 
bluethread; O; DS (509) 


Commelinaceae 
Callisia ornata cD G, : eo scrub 
seling; O; MF (75 
oe erecta L. ane dayflower;O; 
RD (77, 781) 


Cyperaceae 

“Bulbostylis barbata (Rottb.) C.B. Clarke— 
watergrass; O; RD (838) 

Carex gigantea Rudge—giant sedge; C; FF (556, 
661) 

Carex longi Mack.—Lonq's sedge; C; FF (546, 558, 
602, 830) 

Carex verrucosa Muhl.—warty sedge; C; DS (383 

06) 

Cladium jamaicense Crantz—Jamaica swamp 
sawgrass; C; DS coc 

Cyperus croceus Vahl | lge;C;D 
(224) 

Cyperus haspan L.—haspan flatsedge; C; DM 
(571, 539) 

Cyperus odoratus L.—fragrant flatsedge; C; FF 
(545, 774) 

Cyperus polystachyos Rottb.—manyspike 
flatsedge; C; DM (160, 243, 295) 

Cyperus retrorsus Chapm.—pinebarren flatsedge; 
C; DM (136, 247, 293,410 


Ta) 


| 4 


C; DM (104, 232, 294, 437) 

Eleocharis baldwinii (Torr.) Chapm.—Baldwin's 
spikerush; C; DS (423, 430, 876) 

Eleocharis flavescens (Poir.) Urb.—yellow 
spikerush; C; WP (231, 547) 

Eleocharis vivipara Link—viviparous spikerush; C; 


(836) 
Fimbristylis autumnalis (L.) Roem. & Schult.— 
slender fimbry; C; RD (585,878) 
Fimbristylis cymosa R.Br—hurricanegrass;O;DS 
(515) 
Fimbristylis dichotoma (L.) Vahl—forked fimbry; 


*Fimbristylis schoenoides (Retz.) Vahl—ditch 
fimbry; O; RD (851,879, 880) 


BRIT.ORG/SIDA 22(1) 


Fimbristylis puberula (Michx.) Vahl—hairy fimbry; 
O; WP (221, 864) 
Fuirena breviseta (Coville) Coville—saltmarsh 
mbrella-sedge; C; DS (169, 513) 
Fulrena pumila (Torr.) Soreng.—dwarf umbrella- 
sedge; C; DS (233) 
Fulrena scirpoidea Michx.—southern umbrella- 
sedge; C; DS (429) 
Lipocarpha maculata (Michx.) Torr—American 
halfchaff sedge; O; DS (847) 
Rhynchospora cephalantha A. Gray—bunched 
eaksedge; C; FF (521) 
Rhynchospora colorata (L.) H. Pfeiff—starrush 
whitetop; O; WP (111) 
Rhynchospora corniculata (Lam.) A. Gray— 
shortbristle horned beaksedge; O; RD (833) 
ee fascicularis (Michx.) Vah|—fas- 
CDM (122, 135, 182, 255, 


aye 7,485) 


Rhynchospora _fernaldii Gale—Ffernald's 
MF (329 


Rhynchospora inundata (Oakes) Fernald— 
narrowfruit horned beaksedge; C; DS (223, 
5,888 


ote aoe latifolia (Baldwin) W.W.Thomas— 
giant whitetop; O; DS (758) 

Rhynchospora Gals A.Gray—sandyfield 
beaksedge; C; SP (432, 786) 

arpa nee ex A. Gray— 
southern beaksedge; C; DM (831) 

Rhynchospora microcephala (Britton) Britton ex 
Small—bunched beaksedge; C; DS (529, 573) 

Rhyncospora miliacea (Lam.) A. Gray—millet 
beaksedge; C; FF (352, 687) 

Rhynchospora plumosa_ Elliott—plumed 
beaksedge; C; MF (189, 416, 678, 681) 

Scirpus cyperinus (L.) Kunth—woolgrass; O; DM 


Rhynchosnorg micr 
re 


80) 

Scleria baldwinii eh Steud.—Baldwin’s 
nutrush; C; WP (742) 

Scleria reticularis Michx.—netted nutrush; C;WP 
428,859 


<j 


Scleria triglomerata Michx.—tall nutgrass; C; SP 
(187) 
Eriocaulaceae 
Erfocaulon compressum Lam.—flattened 
wort; C; WP (510) 
Bea on decangulare L.—tenangle pipewort; 
O; DS (225, 505) 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 649 


/ bh 
| Vee 
head bogbutton; C; RD (334, 526) 
se eee ala yel- 
low hatpins; C; MF (85, 674, 703) 


f VAFRIPAYY RA ee aia 
Iv VVITIL 


Hecmonbraccae 

achnanthes caroliana (Lam.) Dandy—Carolina 
redroot; Ot MF (200) 

Hydrocharitaceae 

Limnobium spongia (Bosc) Rich. ex Steud.— 
American sponge plant; O; RD (832) 


Hypoxidaceae 
Hypoxis curtissii Rose—common_ yellow 
stargrass, CG gee 


£, 1 
fringed y HOVV SLalUl doo, 


“MF 97, 300, 656) 
lridac 
Iris ae Walter—Dixie iris; C; FF (660) 
Sisyrinchium nashi E.P. Bicknell—Nash’s blue- 
eyed grass; C; MF (668, 729) 
*Sisyrinchium rosulatum E.P. Bicknell—annual 
blue-eyed grass; R; RD (653) 


Juncaceae 

Juncus dichotomus Elliott—forked rush; O; DM 
548, 828) 

Juncus effusus L.—soft rush; C; DS (402,537) 

Juncus marginatus Rostk.—shore rush; C; WP 
(144, 484, 435, 572) 

Juncus megacephalus M.A.Curtis—bighead rush; 


Juncus scirpoides Lar.—needlepod rush; C; DS 
(258, 192, 205, 327,501) 

Liliaceae 

Lilium catesbaei Walter—Catesby’s lily; O; MF; 
threatened (371) 

Melanthiaceae 

Stenanthium densum (Desr.) Zomlefer & Judd— 
crowpoison; R; MF (805) 


es) 


Nartheciaceae 
Aletris lutea Small—yellow colicroot; O; MF (238, 
723) 


Orchidaceae 
Calopogon tuberosus Re Britton et al—tuber- 
ous grasspink; R; 793) 
ea a ees Ane ) Small—Florida butter- 
chid;O;FF;commercially exploited (662) 
ete conopseum R. Br.—green-fly orchid; 
O; FF; commercially exploited (350) 


Habenaria ame Lind|.—toothpetal false 
reinorchid; C; BS (594) 

ae a ecristata Saas Rolfe—giant 

id; O; MF; threatened (81 

Spi ae laciniata (Small) ee 
ladiestresses; O; DS; threatened (764) 

Spiranthes odorata (Nutt.) Lindl.—fragrant 
ladiestresses; O; FF (566) 

Spiranthes praecox (Walter) S. Watson— 
greenvein ladiestresses; O; MF (745) 

Schitr—soldlier’s or- 


xa 


A! 


- - eg: | 
CUATFIE STPUCEUTTIGUCa (L.) 


chid; R; MF (600) 


Poaceae 

Amphicarpum muhlenbergianum (Schult.) 
Hitchc.—blue maidencane; C; BS (427,514 
549, 877,883) 

Andropogon — brachystachyus 
shortspike bluestem; O; MF (563 

Andropogon glomeratus (Walter) Britton et al.var. 
glomeratus—bushy bluestem; O; DS (436) 

Andropogon glomeratus (Walter) Britton et al.var, 
desley ee Cc. Mohr.—purple 

m;O; MF (251,536) 

Pei glomeratus (Walter) Britton et al.var. 
pumilus (Vasey) Vasey ex LH. Dewey-—bushy 
bluestem; F, WF (476) 

Andropogon aa s Ashe—Elliott’s bluestem; C; 
SH (493, 87 

Andropogon ae Michx.—splitbeard 
bluestem; C; SF (469,475) 

Andropogon virginicus L. var. decipiens C.S. 

mpb.—broomsedge bluestem; C; MF 
(471, 486 

Andropogon virginicus L. var. saa Hack.— 
chalky bluestem; O; SF (49 

Aristida palustris (Chapm.) en: 
threeawn; O; MF (845 

Aristida purpurascens Poir. var. tenuispica (Hitchc.) 
Allred—Hillsboro threeawn;C; MF (562,596, 
680) 


Arictidg nirniswra 


Chapm.— 
) 


— 


5 Poir. var. virgata (Trin.) All- 
red—arrowfeather threeawn; O; WP (488 
Aristida ae iciformis Elliott—bottlebrush 

threeawn; C; MF (308, 364, 415, 474) 

Aristi ee stricta Michx. var. beyrichiana ee & 
Rupr.) D.B. Ward—wiregrass; C; MF (407) 
Axonopus furcatus (Fluggé) ee 

carpetgrass; C; RD (801, 829) 
Axonopus fissifolius (Raddi) Kuhlm.—common 


carpetgrass; O;RD (857 


650 


Cenchrus spinifex Cav.—coastal sandbur; C; MF 

Ctenium aromaticum (Walter) AW. Wood— 
toothachegrass; O; MF (519, 886) 

*Cynodon dactylon (L.) Pers. —Bermudagrass; O; 
RD (842) 


*Dactyloctenium aegyptium (L.) Willd. ex Asch. & 
Schweinf—Durban crowfootgrass; C; MF 


(525) 
Dichanthelium commutatum (Schult.) Gould— 
ariable witchgrass; C; MF (401, 605, 861) 
Dichanthelium dichotomum (L.) Gould—cypress 
MF (557, 684) 
Dichanthelium ensifolium (Baldwin ex Elliott) 
ould var. ensifolium—witchgrass; C; WP 
(190, 411,540,675, 712, 865 
Dichanthelium ensifolium (Baldwin ex ae 
d var. unci oe (Trin.) B.F. Hansen & 
Wunderlin— 5 witchgrass;O; MF 7 
Dichanthelium seciel um (Nash) ae & CA. 
Clark—erectleaf witchgrass; C; MF (2 
Dichanthelium laxiflorum (Lam. i 
openflower witchgrass; O; MF (800) 
see) ium ovale (Elliott) Gould & C.A.Clark— 
ggleaf witchgrass; F; MF (717) 
Dichanthelium portoricense (Desv. ex Ham.) B.F. 
Hansen ee 
C; MF (363, 667, 676, 713) 
Digitaria ciliaris (Retz.) Koeler—southern crab- 
grass; O; RD (856, 881) 
eusine indica (L.) Gaertn.—Indian goosegrass; 


‘e) 
= 


eo. 


*E 


—s, 


“Eragrostis atrovirens (Desf) Trin.ex Steud.—Tha- 
lia lovegrass; C; WP (229, 257, 543, 575, 583, 
844) 


Eragrostis elliottil S. Watson—Elliott’s lovegrass; 
» MF (414, 438, 4 
Eragrostis secundiflora J.Presl subsp. oxylepis al 
5.D. Koch—red love-grass; O: MF (81 
Eragrostis virginica (Zuccagni) sit ance 
lovegrass; C; WP (412, 569) 
*Eremochloa ophiuroides (Munro) Hack.— 
centipedegrass; C; RD (872) 
Eustachys petraea (Sw.) Desv.—pinewoods 
fingergrass; C; MF (164, 244 
Leersia hexandra Sw.—southern cutgrass;C; DM 


(542) 
Panicum anceps Michx.—beaked panicum;C;DS 
(252, 137, 406, 835) 


BRIT.ORG/SIDA 22 


_~ 


1) 


Panicum dichotomiflorum Michx.—fall panic- 
grass; O; DM (843) 

Panicum hemitomon Schult.—maidencane; C; 

“Panicum repens L.—torpedograss; C; RD; 
FLEEPC-I (827) 

Pani rm rials 


aidilism PB 


c ex Nees—redtop pani- 
cum; C; BS (439, 490, 848) 
Panicum verrucosum Muhl—warty panicgrass; 
(420) 


atum L.—switchgrass; C; MF (568) 
Paspalum floridanum Michx.—Florida paspalum; 
(246) 


dD : 
CO HCUETT VY 


Paspalum laeve Michaux—field paspalum;O;WP 
84) 
“Paspalum notatum Fluggé var. notatum— 
Bahiagrass; O; MF (186) 
“Paspalum notatum Fluggé var.saurae Parodi— 
Bahiagrass; C; MF (756) 
Paspalum praecox Walter—early paspalum;C; MF 
(226) 
Paspalum setaceum Michx.—thin paspalum; C; 
MF (256, 421, 433, 489, 530, 806) 
“Paspalum urvillei Steud.—Vaseygrass; C; WP 
(245,757) 
*Rhynchelytrum repens (Willd.) C.E. Hubb.—rose 
Natalgrass; C; MF; FLEPPC-II (809) 
Saccharum giganteum (Walter) Pers.—sugar- 
cane plumegrass; C; DS (424, 550, 582) 
*Sacciolepis indica (L.) Chase—Indian cupscale; 
C; DM (862) 
Sacciolepis striata (L.) Nash—American cupscale; 
FF (534) 
Schizachyrium scoparium (Michx.) Nash—little 
luestem; C; MF (492) 
Setaria parviflora (Poir.) Kerguelen—yellow 
PHAGES Gc ve (172,2 vey 


(CW; | | 
i} 


BR 
Ne) 


(9) 
ne) 
a 


IncianarasC MF (391) 

Sphenopholis obtusata (Michx.) Scribn.—prairie 
wedgescale; C; FF (692) 

Sporobolus floridanus Chapm.—Florida 
dropseed;O; MF (431, 863, 887) 

“Sporobolus indicus (L.) R. Br—smutgrass; C; MF 
(434) 

Sporobolus junceus (PBeauv.) Kunth—piney- 
woods saeuiaieaia MF (763) 


pall 


Ponteder 
Pontederia eee L.—pickerelweed;C; BS (109) 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 651 


Smilacaceae 

Smilax auriculata Walter—earleaf greenbrier; C; 
SH (315 

Smilax bona-nox L.—saw greenbrier; C; FF (555, 


Smilax at greenbrier;O;FF (818) 

Smilax feuntelia L aie greenbrier;C; BS (522, 
561,592) 

Smilax pumila Walter—sarsaparilla vine; O; SF 
(689) 


yphaceae 
Typha latifolia _—broadleaf cattail; O; WP (773 


— 


Xyridaceae 

Xyris ambigua Beyr. ex Kunth—coastalplain 
yelloweyed grass; C; MF (165) 

Xyris caroliniana cee yelloweyed 


grass; C; MF (139,814) 
Xyris elliottii Chapm. —Elliott’s y \| yed grass; 
P (133) 


Xyris fimbriata Elliott—fringed yelloweyed grass; 
CG; Ds G89) 

Xyris flabelliformis Chapm.—Savannah yellow- 

rass; O; RD (644) 

*xyris jupical Rich.—Richard’s yelloweyed grass; 
C;RD (145, 166, 544) 

Xyris platylepis Chapm.—tall yelloweyed grass;C; 

(214) 


= 


DICOTS 


Acanthaceae 

Dyschoriste oblongifolia (Michx.) Kuntze— 
oblongleaf twinflower; O; SH (750) 

Ruellia ciliosa Pursh—ciliate wild petunia; O; SH 
(3 


22) 
Adoxaceae 
Sambucus nigra L. subsp. canadensis (L.) Bolli— 
merican elder; O; BS (769) 
Viburnum obovatum Walter—Walter’s viburnum; 
C; FF (346, 686) 


Amaranthaceae 
*Chenopodium ambrosioides L.—Mexican tea;C; 
RD (789) 


Froelichia floridana (Nutt.) Mog.—cottonweed; 
(194) 


Anacardiaceae 

Rhus copallinum L.—winged sumac; O; SH (361) 

Toxicodendron radicans (L.) Kuntze—eastern 
poison ivy; C; DS (533) 


Annonaceae 
Asimina angustifolia Raf—slimleaf pawpaw; O; 
SH (718) 
Asimina obovata oe Nash—big flower paw- 
w; C; SP (650, 704, 785) 


Asimina reticulata ae ex Chapm.—netted 
wpaw; C; SH (449, 636) 

Apiaceae 

Eryngium baldwinii Spreng—Baldwin's eryngo; 
O; FF (87, 737) 

Eryngium yuccifolium Michx.—button rattle- 
snakemaster; O; SH (143 

Oxypolis filiformis (Walter) Britton—water cow- 
bane; C; RD (426, 578 

Ptilimnium capillaceum (Michx.) Rafi_-mock 
bishopsweed; C; DM (105, 759, 790) 


Apocynaceae 

Asclepias feayi Chapm. ex A. Gray—Floricla milk- 
weed; C; MF (78, 96) 

Asclepias humistrata Walter—pinewoodls milk- 
weed: C: SH (696) 

Asclepias longifolia Michx.—longleaf milkweed; 
O;WP (227) 

Asclepias pedicellata Walter—Savannah milk- 

Ww (212) 

Asclepias tomentosa Elliott—velvetleaf milk- 
weed; O; SH 

Asclepias tuberosa L—butterflyweed; O; SH (88) 


fe r 


Aquifoliaceae 

llex cassine L—dahoon,; C; FF (397) 

llex glabra (L.) A.Gray—gallberry;C; DM 
450) 

Araliaceae 

Centella asiatica (L.) Urb.—spadeleaf;C;DS (819) 

Hydrocotyle umbellata L.— flower marsh- 
pennywort; C; DS (654, 721) 

Hydrocotyle verticillata Thunb.—whorled 
marshpennywort; O; FF (804) 

Asteraceae 

Acmella oppositifolia (Lam.) R.K. Jansen var. repens 
(Walter) R.K. Jansen—oppositeleaf spot- 
flawer; O; DM (468) 

Ambrosia artemisiifolia L. 
MEF ( ) 


—_ 


259,362, 


MMon ran ac: 

Baccharis halimifolia _—groundsel tree; C; SH 
(455,477) 

Balduina angustifolia (Pursh) B.L. Rob.— 


652 


coastalplain honeycombhead; GC; SH (208, 
wer) 
Bidens alba (L.) DC. var. radiata (Sch. Bip) R.E. Bal- 
lard ex Melchert—beggarticks; C; RD (285) 
see a Nh subsp. australis L.C 
lee eal rayless goldenrod; C; 

an on 27) 


— 


AARC ASI 
Sper 


EiWiatt Ih 


RD (481) 
Carphephorus corymbosus (Nutt.) Torr. & A. 
Gray—coastalplain chaffhead; O; SF (291) 
Carphephorus odoratissimus (J.F.Gmel.) H. Hebert 
var. aye (DeLaney et al.) Wunderlin 
& B.F. Hansen—vanillaleaf; C; MF (303) 

Po paniculatus (J.F. Gmel.) H. He- 
bert—hairy chaffhead; C; WP (495) 

Chaptalia tomentosa Vent. 
C; DM (580, 611) 

Chrysopsis mariana (L.) Elliott—Maryland 
goldenaster; C; MF (791) 

Chrysopsis subulata Small—scrubland golden- 
aster; C; MF (161, 197, 207, 815) 

Cirsium horridulum Michx.—purple thistle; O; MF 
(691) 

Cirsium nuttallii DC.—Nuttall’'s thistle;O; MF 
203) 


{| | 
VWOUOTTY SUTINOTIT TIE, 


— 


195, 


Conyza canadensis (L.) Cronquist var. pusilla 
(Nutt.) Cronquist—dwarf Canadian 
rseweed; C; RD (171,317, 358) 
— floridana E.B. Smith—Florida tickseed; 
(DS(576) 


creps leavenworthii Torr. & A. Gray 
Leavenworth’'s tickseed; O; MF (158) 
Croptilon ie aie (Nutt.) Raf.—slender 
lane y, i ie alee 
| allot 


= 


5G: SH(I 52, 283, 325) 
Erechtites hieraciifolius (L.) Raf.ex OC.—fireweed; 


Frigeron quercifolius Poir—oakleaf fleabane; C; 
MF (100, 657) 


Erigeron vernus (L.) Torr. & A. Gray—early 
whitetop fleabane; C; MF (162,372,574, 747) 

Eupatorium capillifolium (Lam.) Small ex Porter 
& Britton—dogfennel; C; MF (483 

Eupatorium compositifolium Walter—yankee- 
weed; C; DS (441,458, 478) 

Eupatorium mohrii Greene—Mohr's through- 
wort; C; MF (121, 448, 777) 


BRIT.ORG/SIDA 22(1) 


Eupatorium rotundifolium  L.—roundleaf 
throughwort; C; MF (209, 235) 
Eupatorium serotinum een 
throughwort; CG; DM (34 
Euthamiacaroliniana . ) crane eX ee & Brit- 
r flattop golder C;MF (442) 
ea a pulchella eee rewheeh O; RD (90 
181) 
Gamochaeta antillana (Urb.) Anderb.—Carib- 
bean purple everlasting; O; RD (64 
cama pensylvanica (Willd.) Cabrera— 
vania everlasting; O; RD (641) 
ee a een (W. Bartram) Merr. & F. 
Harper—garberia; O; SP; threatened (512) 
ree pinnatifidum (Schwein. ex Nutt.) 
db.—southeastern sneezeweed; O; DS 
(725) 
Helianthus angustifolius L.—narrowleaf sun- 
flower; CG; DS (464 


on—s 


Helianthus radula (Pursh) Torr. & A. Gray—stiff 
sunflower; O; MF (356) 
Heterotheca subaxillaris (Lam. : Britton & Rusby— 
phorweed; C; SH (366, 444, 535) 


te gronovil eal O; SH (454, 
603) 


Hieracium megacephalon Nash—coastalplain 
hawkweed; C; MF (174, 193, 260, 264) 

lva microcephala Nutt——piedmont marshelder; 
C; DS (466, 531) 

Krigia virginica (L.) Willd.—Virginia dwarfdande- 
lion; C; RD (637) 

Lactuca graminifolia Michx.—grassleaf lettuce;G 
RD (720, 760) 

Liatris gracilis i —slender gayfeather; C; SH 

(250, 375, 392) 

Liatris poo Pursh—fewflower gayfeather; 
O; SH (324) 

Liatris — (L.) Willd. —dense gayfeather;O;SH 
(37 

Liatris wie Nutt.—shortleaf gayfeather; C; 


H (403 

Lygodesmia aphylla (Nutt.) DC.—rose-rush;C; MF 
(81, 302) 

Mikania scandens (L.) Willd.—climbing 


hempvine; O; FF (110, 382, 869) 
Oclemena reticulata (Pursh) G.L.Nesom—white- 
topped aster; C; DS (156, 447, 496, 711) 
Palafoxia intergrifolia (Nutt.) Torr & A. Gray— 
coastalplain palafox; O; SH (154) 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 653 


Phoebanthus grandiflorus (Torr. & A. Gray) S.F. 
Blake—Florida false sunflower; C; MF (129, 


Pityopsis graminifolia (Nutt.) Michx.—narrowleaf 
silkgrass; C; MF (261, 599) 

Pluchea foetida (L).DC.—stinking camphorweed; 
O; DS (343) 

Pluchea rosea R.K.Godfrey—rosy camphorweed; 
C; FF (101, 149, 198, 820) 

ee obtusifolium (L Hilliard & B.L. 
Bur weet everlasting; C; MF (393) 

Pterocaulon oF ee Elliott— 
blackroot; C; MF (79) 

Pyrrhopappus carolinianus (Walter) DC.—Caro- 
lina desertchicory; O; XH (394) 

Sericocarpus tortifolius (Michx.) Nees—whitetop 
aster; C; MF (178, 445, 516) 

Solidago fistulosa Mill—pinebarren goldenrod; 
C; MF (443, 456) 

Solidago odora Aiton var. chapmanii (Torr. & A. 
Gray) ee eeien: goldenrod; C; 
SH (15 

aa stricta Aiton—wand goldenrod; C; MF 

03) 
sept ichum adnatum (Nutt.) G.L.Nesom— 
caleleaf aster; O; MF (597) 

Ee eta carolinianum (Walter) 
Wunderlin & B.F.Hansen—climbing aster; O; 
FF (586) 

pa eet dumosum (L.) G.L.Nesom—rice 

utton aster; C; SH eth ae 

symphyot ae subula (Michx.) 

GL m—annual ok aster; Cc; DM 


+ 
T 
= 


a 
Vernonia gigantea (Walter) Trel. ex Branner & 
Coville—giant ironweed; O; FF (860) 


Betulaceae 

Carpinus caroliniana Walter—American horn- 
beam; O; FF (341) 

Bignoniaceae 

Campsis radicans (L.) Seem.ex Bureau—trumpet 
creeper; O; FF (313 

Brassicaceae 

Lepidium virginicum L.—Virginia pepperweed;C; 
RD (652) 

Cactaceae 

Opuntia humifusa (Raf.) Raf—pricklypear; C; SH 
(217) 


Campanulac 
Lobelia ae L.—cardinalflower;O; FF, threat- 


ned (396) 
ee glandulosa Walter—glade lobelia; O; MF 
(507) 
Lobelia ona Nutt.—white lobelia;C; MF (275, 
722,821) 


Triodanis ae ata (L.) Nieuwl.—clasping 
slo okingglass; O; RD (714) 
aN ees ia marginata (Thunb.) A. DC.— 
southern rockbell; CG; RD (216, 715) 


Caryophyllaceae 

Drymaria cordata (L.) Willd. Ex Schult—West |n- 
dian chickweed; O; RD (607) 

Stipulicida setacea Michx. var. lacerata C.W. 
James—pineland scalypink; C; SP (659) 


Chrysobalanaceae 
Licania michauxii Prance—gopher apple; C; SH 


Cistaceae 

Helianthemum carolinianum (Walter) Michx.— 
Carolina frostweed; C; MF (634, 761) 

Helianthemum corymbosum Michx.—pinebarren 
frostweed; C; SH (180, 616, 639) 

Lechea torreyi (Chapm.) Legg. ex Britton-——pied- 
mont pinweed; C; MF (210,629, 853) 


Clusiaceae 
eee cistifolium Lam.—roundtop St.John’s- 
rt;C; WP (117,213,335, 157,646 

bypecum fasciculatum Lam.—peelbark St. 
John's t; GDS (234, 520) 

beeen. etianoide (L.) Britton et al.— 
pineweeds; C; DS (196, 239, 279, 570) 

nee hypericoides (L.) Crantz—St. 
Andrew’s-cross; C; DS (142, 348, 446, 834) 

Hypericum mutilum L.—dwarf St. John’s-wort; C; 
DS (99, 112) 

Hypericum myrtifolium Lam.—myrtleleaf St. 
John’s-wort; C; WP (86, 177) 

Hypericum reductum (Svenson) W.P. Adams— 
Atlantic St. John’s-wort; O; SH (796) 

Hypericum setosum L—hairy St. John's-wort; O; 
DS (373) 

Hypericum tetrapetalum Lam.—fourpetal St. 
John's-wort; C; MF (176, 206) 


— 


Convolvulaceae 
Ipomoea sagittata Poir—saltmarsh morning- 
glory; O; DS (812) 


Stylisma patens (Desr.) Myint—coastalplain 
dawnflower; O; SH (269, 272) 


— 


Cornaceae 
Cornus foemina Mill—swamp dogwood; O; FF 
(685) 


Nyssa sylvatica Marshall var. biflora (Walter) 
Sarg.—swamp tupelo; C; FF (705, 728, 779) 

Droseraceae 

Drosera capillaris Poir—pink sundew; C;DS (120, 
726) 

Ebenaceae 

Diospyros virginiana L—common persimmon;G 
SF (702) 


Ericaceae 
Bejaria racemosa Vent.—tarflower;G SF (118,292, 
487) 


Ceratiola ericoides Michx.—Florida rosemary; O; 


Gaylussacia dumosa (J. Kenn.) Torr. & A. Gray— 
warf huckleberry; C; SH (669, 738, 795) 
Gaylussacia frondosa (L.) Torr. & A. Gray ex Torr. 

var. tomentosa A. Gray—blue huckleberry;O; 


(649) 
Lyonia ferruginea (Walter) Nutt.—rusty stagger 
pualy O;s F (732) 
Lyonia fruticosa (Michx.) G. : ee 
eer O; SF (612, 733) 
ba ia! ee Lam. es C; DS (287, 560, 
591,6 


Vaccinium re L.—highbush blue- 
berry; O; MF (627) 

Vaccinium myrsinites Lam.—shiny blueberry; C; 
MF (76, 615, 642) 


Vaccinium stamineum L.—deerberry; O; SH (651, 
673) 


Euphorbiaceae 
Acalypha gracilens A. Gray—slender threeseed 
mercury; O; RD (802) 


Chamaesyce hirta (L.) Millsp.—pillpod sandmat; 
O; RD (839) 
Chamaesyce hyssopifolia (L. 
sandmat; C; RD (387) 
Chamaesyce maculata (L.) Small—spotted 
sandmat; O; RD (866) 

Cnidoscolus stimulosus ne Engelm. & A. 
Gray—tread-softly; O; SH (253) 

Croton glandulosus antes conmigo; O; RD 


40) 


Small—hyssopleaf 


BRIT.ORG/SIDA 22(1) 


Croton michauxii G.L. Webster—rushfoil; C; SH 
(131, 788, 816) 
*Phyllanthus urinaria L.—chamber bitter; O; RD 


Stillingia aquatica Chapm.—water toothleaf; O; 
BS (222) 

Stillingia sylvatica L—queen’s delight;O;SH (102 
168) 


Fabaceae 
Amorpha herbacea Walter—clusterspike false 
indigobush; (94 


Apios americana Medik.—groundnut;O;DS (813) 

Centrosema virginianum (L.) Benth.—spurred 
butterfly pea; C; SH (92, 284, 323) 

Chamaecrista fasciculata (Michx.) Greene—par- 
tridge pea; C; SH (163, 278) 

Chamaecrista nictitans (L.) Moench var. aspera 
(Muhl. ex Elliott) H.S. Irwin & Barneby—sen- 
sitive pea; ae SH (321) 

“Crotalaria lanceolata E. Mey—lanceleaf 
rattlebox; O; RD (236, 850) 

*Crotalaria pallida Aiton var. obovata (G. Don) 
Polhill—smooth rattlebox; O; RD (218) 
Crotalaria rotundifolia J.F. Gmel.—rabbitbells; C; 

H (126, 127,326, 405,670 

*Crotalaria spectabilis Roth—showy rattlebox; C; 
RD (479) 

*Desmodium incanum DC.—tickfoil; C; RD (248, 
386) 


Desmodium paniculatum (L.) DC.—panicled 
ticktrefoil; O; SH (408, 470) 

*Desmodium triflorum (L.) DC.—threeflower 
ticktrefoil; O; 07) 

Galactia elliottii Nutt.—Elliott’s milkpea; O; SH 


@ 


Galactia regularis (L.) Britton et al—eastern 
milkpea; O; MF (824) 

Galactia volubilis (L.) Britton—downy milkpea;C; 
SH (357, 767, 784, 854) 

Indigofera voc Mill—Carolina indigo; C; 


ese 


*In fera hirsuta re neice Cc: ae Se 
(826) 

Lespedeza hirta (L.) Hornem.—hairy lespedeza; 
C; SF (891) 


Lupinus diffusus Nutt.—skyblue lupine; O; SH 
(635) 


*Macroptilium lathyroides (L.) Urbo.—wild 
bushbean; O; RD (124) 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 655 


*Medicago lupulina L—black medick;O;RD (710) 

Mimosa quadrivalvis L. var. angustata (Torr. & A. 
Gray) Barneby—sensitive brier; O; SH (268) 

Rhynchosia michauxii Vail—Michaux's snout- 
bean; O; SH (671) 

Senna obtusifolia (L.) H.S. Irwin & Barneby— 
coffeeweed; O; RD (868 

Tephrosia chrysophylla Pursh—scurf hoarypea;O; 
RD (867) 

Tephrosia hispidula (Michx.) Pers—sprawling 
hoarypea; O; SH (885 

Vicia acutifolia Elliott—fourleaf vetch;O; FF (708) 

Fagaceae 

Quercus chapmanii Sarg.—Chapman's oak; O; SF 
(695, 701) 

Quercus geminata Small—sand live oak; C; SH 

53) 

Quercus incana W.Bartram—bluejack oak; O; SH 
(694) 

Quercus laevis Walter—turkey oak; C; SH (472, 


473) 
Quercus laurifolia Michx.—laurel oak; O; FF (314 


3) 
Quercus margaretta Ashe ex Small—sand post 
oak; R; SH (762) 
Quercus minima (Sarg.) Small—dwarf live oak;C; 
SH (179, 320, 419) 
Quercus myrtifolia Willd —myrtle oak; C; SF (333, 
462) 


Quercus nigra L.—water oak; O; MF (309) 
Quercus virginiana Mill.—live oak; O; DS (746) 


Gelsemiaceae 

Gelsemium sempervirens (L.) Aiton f—yellow jes- 
samine; O; SF (620) 

Gentianaceae 

Sabatia brevifolia Raf—shortleaf rosegentian;O; 
MF (286, 301) 

Sabatia calycina (Lam.) A. Heller—coastal 
rasegentian; O; FF (739) 

Sabatia grandiflora (A.Gray) Small—largeflower 
rosegentian; C; DM (262, 390, 518) 


Geraniaceae 

Geranium caro 
O; RD (709) 

Haloragaceae 

Proserpinaca pectinata Lam.—combleaf 
mermaidweed; O; DM (564, 741, 768) 


inianum L.—Carolina cranesbill; 


Iteaceae 
Itea virginica L—Virginia willow; O; BS (664) 
Lamiaceae 
Callicarpa americana L—American beautyberry; 
> MF (119) 
Hyptis alata (Raf.) Shinners—musky mint; C; DM 
(274, 331, 345) 
*Hyptis are is (Rich.) 
bushmint; O; FF (173, 276) 
Lycopus rubellus Moench—taperleaf water- 
rehound; F;DM (511) 
Piloblephis rigida (W. Bartram ex Benth.) Raf— 
wild pennyroyal; O; MF (63 
a alvia eae O; BS (606) 
mall—Florida scrub skull- 


Brig.—tropical 


larin nreni 


cap; C; SH (132) 
Teucrium canadense L.—woodsage;C; DM (113) 
Trichostema dichotomum L.—forked bluecurls;C; 


MF (377) 


Lauraceae 

*Cinnamomum camphora (L.) J. 
C hortree; R; BS; FLEPPC-I (734) 

Persea palustris (Raf.) Sarg—swamp bay; C; FF 

(289, 353) 


Pres|— 


Lentibulariaceae 

Pinguicula pumila Michx.—small butterwort; O; 
MF (588) 

Utricularia foliosa L_—leafy bladderwort; O; DS 


O; BS (626) 
Utricularia juncea Vah|—s 


ba | 


wthern bladderwort: 
Utricularia subulata L.—zigzag bladderwort; O; 


Linaceae 

Linum medium (Planch.) Britton var. texanum 
(Planch.) Fernald—stiff yellow flax; O; WP 
(797) 


Logan 

oor eee (.F. Gmel.) Torr. & A. Gray—lax 
or 97) 

Mitreola sessilifolia (J.F. Gmel.) G. Don—swamp 
ornpod; C; WP (148, 370, 379, 852) 

Lythra 


ae nee (Jacq.) J.F. Macbr.—Co- 
lombian waxweed; C; FF (219, 508, 799) 


656 


Lythrum alatum Pursh var. lanceolatum (Elliott) 
rr.& A. Gray ex Rothr.—winged loosestrife; 
O; MF (277) 
Magnoliaceae 
Magnolia virginiana L.—sweetbay; R; DS (658) 


Malvaceae 
*Urena lobata L.—caesarweed; C; FF; FLEPPC-I| 
(500) 


Melastomataceae 

Rhexia cubensis Griseb.—West Indian meadow- 
beauty; O; DS (183 

Rhexia mariana L._—pale meadowbeauty; C; DS 
(84, 697 

Rhexia nuttallii CW. James—Nuttall’s meadow- 
beauty; O; MF (185, 506) 


Myricaceae 
Myrica cerifera L—wax myrtle; C; MF (332, 693) 


Nymphaeaceae 
Nymphaea odorata Aiton—American white 
WatenINys O; ae 2) 

4 Aj 


Nuphar UGVETIC VANILOTT) F | 
BS (754) 

Olacaceae 

Ximenia americana L.—tallow wood; O; SP (460) 


eaceae 
Fraxinus caroliniana Mill.—Carolina ash; C; FF 
(107, 337, 778) 


Onagraceae 
Gaura angustifolia Michx.—southern beeblos- 
m; O; MF (125, 128) 
pce linearis Walter—narrowleaf prim- 
willow; C; DS (280, 385,451) 

eee linifolia Poir.—southeastern prim- 
rosewillow; C; DM (265, 699) 

Ludwigia maritima R.M. Harper—seaside prim- 
rosewillow; O; MF (147, 201, 263) 

Ludwigia dial Michx.—smallfruit prim- 
rosewillow; O; DS (297) 

Ludwigia ets (Jacq.) Raven—Mexican 
primrosewillow; O; DM (384, 541) 

“Ludwigia peruviana (L.) H. Hara—Peruvian 
primrosewillow; O; RD (825) 

Ludwigia repens J.R. Forst.—creeping prim- 
rosewillow; C; FF (772) 

Ludwigia suffruticosa Walter—shrubby prim- 
rosewillow; O; DS (240) 


BRIT.ORG/SIDA 22(1) 


Oenothera laciniata Hill—cutleaf eveningprim- 
se; O; RD (204, 638) 


Orobanchaceae 

Agalinis fasciculata (Elliott) Raf —beach false fox- 
glove; C; MF (359, 381) 

Agalinis linifolia (Nutt.) Britton—flaxleaf false 
floxglove; O; DM (846) 

Agalinis tenuifolia (Vahl) Raf—slenderleaf false 

-O; MF (502) 

Buchnera americana L.—American bluehearts;O; 
MF (123, 755) 

ee cassioides (J.F. Gmel.) S.F. Blake 
yaupon blacksenna; O; MF (237,376 422) 

ie pectinata Pursh—piedmont black- 
senna; O; SH (368) 


— 


x< 
‘Q 


Oxalidaceae 
Oxalis corniculata L—common yellow wood- 
sorrel; O; RD (617) 


Passifloraceae 
Passiflora incarnata L_—purple passionflower;O; 
D (155) 
Plantaginaceae 
Plantago virginica L.—Virginia plantain; O; RD 
(648) 


Polygalaceae 

Polygala cymosa Walter—tall pinebarren milk- 
wort; O; DS (724) 

Polygala lutea L—orange milkwort; CG; MF (83, 
188) 

Polygala nana (Michx.) DC._—candyroot;C; MF 


Polygala rugelii Shuttlew. ex Chapm.—yellow 
milkwort;O;WP (175 
/ i) NAA ta lAlai rie 


pe) 


C; MF (98, 211, 748) 
Polygala violacea Aubl.—shov y milkwot t;C; MF 
(93) 


Polygonaceae 
Polygonella gracilis Meisn.—tall jointweed; O; SH 
(367, 404, 567) 

Polygonella polygama (Vent.) Engelm. & A. 
Gray—October flower; C; SH (395, 461) 
Polygonum hydropiperoides Michx.—swamp 

smartweed; C; DS (114, 399, 482) 
Polygonum punctatum Elliott—dotted smart- 
weed; C; RD (400) 


Rumex hastatulus Baldwin—heartwing dock; O; 
RD (719) 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 657 


Portulacaceae 


*Portulaca amilis Soeg.—Paraguayan purslane; 
O; RD (8 


08) 
Primulaceae 
Samolus valerandi L. subsp. parviflorus (Raf. 
Hulten—pineland pimpernel; O; FF (349) 


a 


Rhamnaceae 
Berchemia scandens (Hill) K. Koch.—rattan vine; 
O; FF (336) 


Rosaceae 

Photinia pyrifolia (Lam.) K.R. Robertson & J.B. 
Phipps—red chokecherry; O; DS (628) 

Prunus caroliniana (Mill.) Aiton—Carolina 
laurelcherry; O; RD (631) 

Prunus serotina Ehrh.—black cherry; O; RD (632) 

Rubus argutus Link—sawtooth blackberry;C; DS 
(645, 663) 


Sapindaceae 

Acer rubrum L.—red maple; C; FF (590) 

Saururaceae 

Saururus cernuus L.—lizard’s tail; C; FF (585) 

Solanaceae 

Solanum americanum Mill.—American black 
nightshade; O; FF (593) 

Tetrachondraceae 

Polypremum procumbens L.—rustweed; C; RD 
(130) 


Theaceae 
Gordonia lasianthus (L.) J. Ellis—loblolly bay; O; 
BS (288, 559) 
Ulmaceae 
Ulmus alata Michx.—winged elm; O; FF (682) 
Ulmus americana L.—American elm; O; FF (619 


Rubus cuneifolius Pursh—sand blackberry:C:MF 35) 
(730) ah Ulelcaceas 
uthern dewberry;O; FF ylindrica (L.) Sw.—false nettle; C; DS 
(792) (342, 584) 


biaceae 
Cephalanthus occidentalis L—common button- 
bush; C; DS (524, 770) 
Diodia teres Walter—rough buttonweed; C; SH 
(304) 
Diodia virginiana L—Virginia buttonweed;C;BS 
(106, 199, 388) 
Galium tinctorium L.—stiff marsh bedstraw;C; RD 
(598) 
Houstonia ee tie (J.F. Gmel.) Standl.—in- 
e; C; MF (589, 604) 
Mee ay repens L.—partridgeberry; O; FF 
688) 


— 


340, 


Oldenlandia uniflora L.—clustered mille graine; 
C; DS (413, 882 — ) 
Psychotria nervosa wild coffee; C; FF (339) 
a sulzneri ie coffee; 
FF (338 


ey Sie +. coal AA 
clover; CG; RD (202) 

Spermacoce assurgens Ruiz & Pav.—woodland 
false buttonweed; O;DM (241) 

ean prostrata Aubl.—prostrate false 
buttonweed; O; FF (299) 


Salicaceae 
Salix caroliniana Michx.—Carolina willow; C; RD 
(633) 


Verbena 
Phyla ae nee (L.) Greene—turkey tangle 
fogfruit; C; RD (89) 
Bae brag iensis Vell—Brazilian vervain; O; 
D (775) 


ie scabra Vah|—sandpaper vervain; O; MF 
(849) 

Veronicaceae 

Bacopa monnieri (L.) Pennell—herb-of-qrace; O; 
DM (625 


Gratiola hispida (Benth. ex Lindl.) Pollard-——rough 
gehyssop; O; MF (140, 141) 
Gratiola pilosa Michx.—shaggy hedgehyssop;O; 
B , 167, 766) 


Gratiola ramosa Walter—branched hedgehys- 
sop; O; DS (698) 


Linaria canadensis (L.) Chaz —Canadian toadflax; 
C; DS (622, 643) 

*Lindernia crustacea (L.) F. Muell—M 
false pimpernel; O; RD (440) 

Mecardonia acuminata (Walter) Small subsp. 
peninsularis (Pennell) Rossow—axilflower;C; 
WP (230, 743) 

Micranthemum glomeratum = (Chapm.) 

ners—manatee mudflower;O;WP (623) 
ee dulcis L—sweetbroom; C; MF (151) 


alaysian 


Violaceae 

Viola lanceolata L—bog white violet; O;DS (624) 
Viola palmata L.—early blue violet; O; MF (647) 
Viola primulifolia L.—primroseleaf violet; C; DS 


BRIT.ORG/SIDA 22(1) 


Vitaceae 
Ampelopsis arborea (L.) Koehne—peppervine;O; 
50) 


Parthenocissus quinquefolia (L.) Planch.—Virginia 
) 


Viola sororia Willd—common blue violet;O;FF Vitis rotundifolia Michx. == CeIn) C;MF (360) 
21) Vitis shuttleworthi House -O;MF 


(736) 


Viscaceae 
Phoradendron leucarpum (Raf.) Reveal & M.C. 
Johnst.—oak mistletoe; O; BS (465) 


ACKNOWLEDGMENTS 


We are grateful to Darrell Freeman and Eugene Kelly of the Southwest Florida 
Management District for providing collection permits, access to the property, 
and other kinds of assistance. We thank Doug Coleman and John Kunzer for 
assistance in the field. We also thank Bruce F Hansen for assistance in plant 
identification and for his helpful comments on the manuscript and Kathleen 
Hotchkiss for assistance with graphics. 


REFERENCES 


CHEN, E.and J.F.Gereer. 1990. Climate. In: R.L. Myers and J.J.Ewel,eds. Ecosystems of Florida. 
University Presses of Florida, Gainesville. Pp. 11-34. 

Coie, N.C.and M.A. Gartano. 2003. Notes on Florida’s endangered and threatened plants. 
Florida Department of Agriculture and Consumer Services, Gainesville. 

FLorIDA Exotic Pest PLANT Council (FLEPPC). 2005. List of Florida's invasive species. Florida 
Exotic Pest Plant Council. (http: 

FLoriIDA NaturaAt AREAS INveNToRY (FNAI). 
Tallahassee. 

FLORIDA NATURAL AREAS INVENTORY (FNAI). 2004. Natural community mapping project of Starkey 
Wilderness Preserve, Southwest Florida Water Management District. Florida Natural 
Areas Inventory, Tallahassee. 

Lawson, S.F., R.P. INcatts, and C. Bayess. 1981. Preserving for the future: A history of the J.B. 
Starkey Wilderness Park. Department of History, University of South Florida, Tampa. 
Reprinted by Southwest Florida Water Management District, Brooksville. 

Mitanicu, J.T. 1994, Archaeology of Precolumbian Florida. University Press of Florida, 
Gainesville. 

Scott, 1.M, K.M. CAMPBELL, F.R. Rupert, J.D. ArtHur, T.M. Missimer, JM. Ltoyp, JW. Yon, and J.G. Dun- 
cAN. 2001. Geologic map of the state of Florida. Florida Geological Survey in coopera- 
tion with the Florida Department of Environmental Protection.U.S. Department of the 
Interior, U.S. Geological Survey, Center for Coastal Geology, Washington. 

SouTHwest Florina Water MANAGEMENT District (SWFWMD).1988.Ground-water resource avail- 
ability inventory: Pasco County, Florida. Brooksville 


VV ww.fleppc.org). 
1990. Guide to the natural communities of Florida, 


FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 659 


SoutHwest Floriba WaTER MANAGEMENT District (SWFWMD). 1990.A plan for the use and man- 
agement of the Starkey tract. Brooksville 

Sankey, D.L. 1982. Soil survey of Pasco County, Florida. U.S. Department of Agriculture, Soil 
Conservation Service in cooperation with University of Florida, Institute of Food and 
Agriculture Sciences Department and Florida Department of Agriculture and Con- 
sumer Services. U.S. Government Printing Office, Washington. 

United States Geotocicat Survey (USGS). 1974.Odessa Quadrangle, Florida 7.5 Minute Series 
(topographic). United States Geological Survey, Department of the Interior, Washington 

Weis, M.2004. Pasco county property appraiser. Dade City. (http://appraiser.pascogov.com/). 

WETTERHALL. W.S. 1964. Geohydrologic reconnaissance of Pasco and southern Hernando 
Counties, Florida. Florida Geological Survey Report. Florida Geological Survey, Talla- 
hassee. 

Witey,G.R.1949. Archeology of the Florida Gulf Coast. Smithsonian Miscellaneous Collec- 
tions, 113:1-559. Smithsonian Institute, Washington. 

Wunoerun, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, 
Gainesville. 

WunNDERLIN, R.P.and B.F. HANseN. 2003. Guide to the vascular plants of Florida, 2nd ed. Univer- 
sity Press of Florida, Gainesville. 

Wunpbertin, R.P. and B.F. Hansen. 2005. Atlas of Florida vascular plants [S.M. Landry and K.N. 
Campbell (application development), Florida Center for Community Design and Re- 
search] Institute for Systematic Botany, University of South Florida, Tampa. (http:// 
www.plantatlas.usfedu). 


BRIT.ORG/SIDA 22(1) 


Book NOTICE 


JULIAN A. STEYERMARK, PAUL E. Berry, Kay YatskiEvycH, and Bruce K. Horst (Eds.). 
Flora of the Venezuelan Guayana. Vol. 9, Rutaceae-Zygophyllaceae. (ISBN 
1-930723-47-4, hbk.). Missouri Botanical Garden Press, PO. Box.299, Saint 
Louis, MO 63166-0299, U.S.A. (Orders: http://www.mbgpress. EOE Oe 608 
pp., 502 b/w line drawings, 71/4" x 10 1/2". 

The final volume of the series, concluding the taxonomic treatment of all native and naturalized 

vascular plant families known to occur in the Venezuelan Guayana. Forty-six contributors in Vol- 

ume a treat 48 uli oe eeners and 971 species lystrated es he figures. C eratophyllaceae is 


(from Theagens: Muntingiaceae (from Tiliaceae), oF Semaine (from Simaroubacene)~ —Guy 
Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, 
Editor’s note—Two new names are published: Sterculia kayae PE. Berry, nom. nov. a Xyris 


sulcatifolia Kral, nom. nov. 


SIDA 22(1): 660. 2006 


VASCULAR FLORAS OF SONOITA CREEK STATE NATURAL 
AREA AND SAN RAFAEL STATE PARK: ARIZONA'S FIRST 
NATURAL-AREA PARKS 


Steven P. McLaughlin 


Offi FA id | q eoales 
ay ed at 
THE GEIIVE 


Tucson, Aaa 85721 U S A. 
spmcl@ag.arizona.edu 


ABSTRACT 


J CRC - ft 


Sonoita Creek State N 1 Area (SCSNA) and San Rafael State P two natural- 
area parks in Arizona established with funding provided by the Heritage Initiative. Both sites are in 
Santa Cruz County, and both have riparian communities that are rare in the arid Southwest. The 
upland vegetation of SCSNA is primarily semidesert grassland, while that of SRSP is Plains grass- 
land. The flora of SCSNA includes 525 native species and 36 exotic species; that of SRSP includes 407 


native species and 50 Significant taxa include one new species, Phacelia sonoitensis, 
found at SCSNA. Taxa new to fie flora of Arizona include Pennellia robinsonii Pa Aes 
minima (SCSNA), and Sibara virginica (SRSP). Lilaeopsis schaffneriana ssp. recury 


species, was found at SRSP. The ies of both SCSNA and SRSP are strongly eee in their ee 
ties. Species found at SCSNA but not at SRSP have relatively greater affinities with the flora of the 
Sonoran Floristic Province, while those ae at SRSP but not SCSNA have a greater affinity with the 
Great Plains. 


RESUMEN 


Sonoita Creek state Nancie ance SONA) y San Rafael State Park (SRSP) son los primeros parques 
aaa ge Herencia. Ambos lugares estan en el 
ondado de Santa Cruz, i id raras en el Suroeste arido. La 
vegeta del aaeao de SCSNA es un pastizal semidesértico y la de SRSP es un pastizal de las 
andes Llanuras. La flora de SCSNA ee 525 pees taclesnes y 36 a sce exoticas; i de 
incluye 407 especies indigenas y 50 
una especies nueva, pagers noite nsis, s encontrada € en eae Entre De especies nuevas para la 
flora de Arizona se incluye ), y Sibara 
virginica (de SRSP). ent schaffneriana ssp. recurva, una especie en peligro fue aren en 
SRSP. Las floras de SCSNA y SRSP son fuertemente Apachianas en sus afinidades floristicas. Las 
especies encontradas en SCSNA pero no en SRSP tienen una afinidad mas grande con la flora de la 
provincia floristica de Sonora y las especies encontradas en SRSP pero no en SCSNA tienen una 
uras. 


afinidad mas grande con las Grandes Llanur 

INTRODUCTION 
Arizona voters passed the Heritage Initiative by a nearly two-thirds margin in 
the November 1990 general election. The Heritage Initiative allocates funds from 
the state lottery to Arizona State Parks and Arizona Game and Fish Depart- 
ment. Part of the funds for Arizona State Parks are designated for the acquisi- 


SIDA 22(1): 661 ~ 704, 2006 


662 BRIT.ORG/SIDA 22(1) 


tion of “natural areas,” defined in the initiative as “parcels of land or water that 
contain examples of unique natural terrestrial or aquatic ecosystems, rare spe- 
cies of plants and animals, or outstanding geologic or hydrologic features.” 

The first two acquisitions of large acreage for natural areas were both in 
Santa Cruz County, in southern Arizona, adjacent to the Mexican state of Sonora. 
The Sonoita Creek State Natural Area (SCSNA) in central Santa Cruz County 
was purchased in January 1994 for $2.8 million. The natural area feature justi- 
fying this purchase was Sonoita Creek and its associated gallery forest. Sonoita 
Creek is one of the few perennial or near perennial streams in southern Ari- 
zona. The SCSNA purchase included 4.5 km of perennial stream, along with 
1,990 ha of watershed north of the creek, including intermittent wetlands in 
the streams of Fresno Canyon and Coal Mine Canyon. Sonoita Creek, Fresno 
Canyon, and Coal Mine Canyon provide habitat for the Gila topminnow 
(Poeciliopsis occidentalis occidentalis), a federally listed endangered species of 
fish. 

The second acquisition was San Rafael State Park (SRSP) in extreme south- 
eastern Santa Cruz County. The 8,900 ha San Rafael Land and Cattle Company, 
which included the San Rafael de la Zanja land grant, was purchased by the 
Nature Conservancy in December 1998 for $1] million. Arizona State Parks pur- 
chased the 1,440 ha of patented land along the Arizona-Sonora border along 
with a conservation easement on the entire ranch from the Nature Conservancy 
for $8.6 million in January 1999. Natural-area values cited for this purchase in- 
cluded the unique grassland communities of the San Rafael Valley and signifi- 
cant riparian and wetland habitat along the upper Santa Cruz River and at Sharp 
Spring and Heron Spring. The wetlands provide critical habitat for at least two 
endangered species, the Huachuca water umbel (Lilaeopsis schaffneriana ssp. 
recurva) and Gila topminnow. 

Most of the watershed adjacent to SCSNA is mapped as semidesert grass- 
land (Brown 1994), which is an extensive area in Arizona and adjacent New 
Mexico, Sonora, and Chihuahua varying in physiognomy and species compo- 
sition. Grasses generally are not the dominant lifeform; instead there is a com- 
plex mixture of shrubs, small trees, succulents, rosette plants, and subshrubs, 
with the exact physiognomy and species composition determined by drought, 
the seasonal distribution of precipitation, periodic freezes, soils, grazing his- 
tory, and fire (Burgess 1995). Burgess suggested the term “Apacherian mixed 
shrub savanna” for this complex mosaic of subtropical, semiarid vegetation 
found above desert and below oak savannas. In contrast, the San Rafael Valley 
is mapped as “Plains grassland” (Brown 1994) and is dominated by perennial 
bunchgrasses and oak savannas with very few leguminous trees and shrubs, 
subshrubs, cacti, or rosette succulents. 

The primary objective of this study was to inventory the floras of these 
two natural area parks. The study also provided an opportunity to address a 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 663 


second objective, to contrast the floras of semidesert grassland and Plains grass- 
lands in southeastern Arizona. 

STUDY AREAS 
Table 1 lists the latitude, longitude, area, elevational relief, mean annual pre- 
cipitation, and mean January and July temperatures for both SCSNA and SRSP. 
Below are brief descriptions of the major physical features, soils, vegetation, 
and recent history of each park. 


Sonoita Creek State Natural Area 

The SCSNA is located along and north of Sonoita Creek downstream from 
Patagonia Lake State Park. It isan area of rugged terrain with moderate to steep, 
rocky slopes. Fresno Canyon hasan intermittent stream and extends from north 
to south across the center of the park. Coal Mine Canyon is an east-west trend- 
ing tributary to Fresno Canyon; the north-facing slopes of Coal Mine Canyon 
provide the most mesic terrestrial habitats within the park. A spring in the 
northeast corner of the park appears to be permanently wet. Elevations range 
from 1,080 m near the junction of Sonoita Creek and Fresno Canyon in the south- 
west corner of the park, to 1310 m at a point near the northern boundary. Fig- 
ure 1 includes a map of SCSNA showing the major drainages, representative 
upland and riparian habitats are shown in Figure 2. 

The exposed surface of the uplands adjacent to Sonoita Creek is that of the 
relatively impervious Nogales Formation, formed by the deposition of fine to 
coarse-grained alluvial fan deposits derived from pre-existing volcanic rocks. 
These deposits have been subject to subsequent folding and erosion, creating a 
complex topography of incised drainages and slopes of all aspects. 

Upland soils at SCSNA include: (1) Graham soils, mostly gravelly or cobbly 
loams on more gentle slopes at lower elevations in canyons, usually at the base 
of tuffaceous mountains; (2) Lampshire-Graham-Rock Outcrop associations, 
shallow, cobbly loams on steeper slopes, and (3) Rock Outcrop-Lithic Haplustolls, 
cobbly loams found in the dissected terrain in the southeast corner of the park 
(Richardson et al. 1979). The soils on the flood plain of Sonoita Creek are deep, 
well-drained sandy loams of the Comoro Association. 

The gallery forest along Sonoita Creek is dominated by cottonwood 
(Populus fremontii), ash (Fraxinus velutina), willows (Salix gooddingii and S. 
bonplandiana), and walnut (JJuglans major). On terraces adjacent to the gallery 
forest there is a dense bosque of mesquite (Prosopis velutina), hackberry (Celtis 
reticulata), elderberry (Sambucus mexicana), and one large colony of tree of 

eaven (Ailanthus altissima) 

The vegetation of the upland portions of SCSNA is mostly semidesert grass- 
land, with small sections of oak (Quercus emoryi and Q. oblongifolia) savanna, 
mostly confined to the north-facing slope of Coal Mine Canyon. Species com- 
position of the semidesert grassland is highly variable. The most common plants 


664 BRIT.ORG/SIDA 22(1) 


Taste 1. Physical features of Sonoita Creek State Natural Area (SCSNA) and San Rafael State Park 
SRSP). Climate data are from Sellers et al. (1985); representative stations used are Nogales 6 N (31° 
26'N,110° 55'W, 1160 m elev.) for SCSNA and San Rafael Ranch (31° 21' N, 110° 37' W, 1446 m elev) 
for SRSP. 


Feature SCSNA SRSP 
Latitude 31° 30'N 31° 21'N 
Longitude 110° 54'W 110° 36'W 
Area (ha) 1990 1440 
Minimum elevation (m) 1080 1395 
Maximum elevation (m) 1310 1475 
Elevational relief (m) 230 80 

Mean annual precipitation (mm) 430 440 

Mean January temperate (°C) 74 5.6 

Mean July temperature (°C) 25.9 23.5 


include mesquite, hopbush (Dodonaea viscosa), coral-bean (Erythrina 
flabelliformis), beargrass (Nolina microcarpa), sotol (Dasylirion wheeleri), 
yucca (Yucca thornberi), shindagger (Agave schottii), acacia (Acacia greggii), 
mimosa (Mimosa aculeaticarpa var. biuncifera), and prickly pears (Opuntia 
spp.). 

Prior to acquisition the site was primarily used for cattle grazing: trespass 
cattle were frequently observed during the floristic inventory. There is also evi- 
dence of historic small-scale mining activity on the site. The New Mexico and 
Arizona Railroad was constructed along Sonoita Creek during 1881-1882. Its 
bridges washed out on many occasions, and after a particularly devastating 
flood in 1929 the section of the line along Sonoita Creek was abandoned (Myrick 
1975). Sections of the railroad bed are evident along most of the reach of the 
stream, usually at the edge of the floodplain. 


San Rafael State Park 

The SRSP is located in the San Rafael Valley, an isolated valley bounded on the 
west by the Patagonia Mountains, on the north by the Canelo Hills, and on the 
east by the Huachuca Mountains. SRSP occurs at the southern end of the valley 
adjacent to the US-Mexico international border. The San Rafael Valley is the 
headwaters of the Santa Cruz River, which runs north to south down the cen- 
ter of the valley. During the period of field work for this study the Santa Cruz 
River ran continuously where it crosses the SRSP, supporting a variety of wet- 
land plant communities. Sharp Spring and Heron Spring are water courses flow- 
ing northeast to southwest into the Santa Cruz River: both have intermittent 
pools and support communities of wetland plants. Parker Canyon wash is a 
major drainage just entering SRSP in its southeastern corner. Figure lincludesa 
map of SRSP showing the major drainages; representative upland and riparian 
habitats are shown in Figure 3. 


RAI APFEA AND SAN RAFAFI STATE PARK 665 


DCCVY NATII 


MCLAUGHLIN, FLORA OF SONOITA 


Sonoita Creek State Natural Area Morning Star Ranch 
i 


ov 


7 spring 
? c 
. c 
§ Hl A : 
‘\ Chivas Sige Sf ey 
*y, Tank r Cons 4 
i : Mj 
? \ ne Co 
§/ “om, on 
Rio Rico X . oS | 
Subdivision — | ) ss 
; - ele 
: QA 7 
* we Ss 
2 


1000 0 1000 2000 Meters 
— _ Park Headquarters 
(Historic Greene Ra ol 


k, Santa Cruz C ty, Arizona 


RA 
Fic. 1. Map 


Topography on the park consists mostly of low, gently sloping hills and 
shallow drainages. There are no bedrock outcrops. The Santa Cruz River is cur- 
rently entrenched about 2 to 4 m within its floodplain, which averages about 
0.5 km in width. 

Soils along the floodplains of the Santa Cruz River, Sharp Spring, and Heron 
Spring are all mapped as Guest Soils, deep loams to clay loams (Richardson et 
al. 1979). The peepee soils of Parker Canyon are gravelly sandy loams of the 
Grabe-Com plex. Most of the upland areas have gentle slopes with grav- 
elly clay loans of the Bernardino-Hathaway Association, clay loams of the 

White House Association, or deep clay loams of the Pima Association. In the 
extreme southeast corner of SRSP on the south side of Parker Canyon, there are 
calcareous gravelly sandy loams of the Hathaway Association on slopes of 20 

to 50 percent steepness. The high clay content of most of the soils of the San 
Rafael Valley is probably associated with their resilience to high grazing pres- 
sure and resistance to invasion by shrubs and cacti. 

The vegetation of the San Rafael Valley is primarily “Plains grassland”, a 
true grassland dominated by grama grasses (Bouteloua spp.), wolftail (Lycurus 
setosus), cane bluestem (Bothriochloa barbinodis), Plains lovegrass (Eragrostis 


ina), both on flood- 


bo 
an RY W 
TIQHIt], Yanler y 


£ 


(Cl 


) 


Pre? 


(lower right). 


Fic 7 epee C¢ * tf a ge ee 
: 
Int Tene parse pea? soph a 
I I PI 
(Salix spp.) al S ita Creek (I 
Pi g 


999 


L)@Z VaIS/OYO'LIYS 


10 VdO1d ‘NIHONV TDW 


MUVd JLVLS 1UVIVd NVS ONY Vad 


Fie 2 Tha be seit, +¢ l C4#a4 I 1 P| £C p6f TV. +h a | £D. L Lin Hy { 
* re 
- +) i fc Di a Se +4] IT ba \ kL bo x. j* . eae } ony . 
} y ) p ( pp.) adj | ghtii), cottonwood, and willow on 
fi Pe f1 1 £cClL L *1 I Dia I Md fl * L \ 
Pv \ t \ TIQHU). <) 


668 BRIT.ORG/SIDA 22(1) 


intermedia), Texas bluestem (Schizachyrium cirratum), spider grass (Aristida 
ternipes), and other perennial bunchgrasses. Broadleaf annual and perennial 
herbs are an important component of the vegetation, particularly species in 
the families Euphorbiaceae, Asteraceae, and Fabaceae. The abundance of non- 
grass components in the vegetation is likely due to intensive grazing in the last 
150 years. Oak savannas are present in the northwestern portion of the park 
and along Parker Canyon wash. 

There is an intermittent gallery forest along the Santa Cruz River, Sharp 
Spring, and Heron Spring composed mostly of cottonwood and willows (Salix 
gooddingii, S. lasiolepis, and S. exilifolia). Herbaceous vegetation along the Santa 
Cruz River is dominated by spikerushes (Eleocharis spp.) and true rushes 
(Juncus spp.). Grazing pressure in the wetlands along the Santa Cruz River, Sharp 
Spring, and Heron Spring appears to have been particularly heavy, and exotic 
species are locally abundant in these areas, particularly sweet clover (Melilotus 
officinalis), meadow fescue (Festuca arundinacea),and Kentucky blue grass (Poa 
pratensis). Most of the floodplain of the Santa Cruz River within SRSP was not 
cultivated, and this area contains an excellent example of sacaton (Sporobolus 
wrightii) grassland, a community that has disappeared from much of the South- 
west due to lowering of water tables. 

Hadley and Sheridan (1995) provided a detailed history of land use in the 
San Rafael Valley. The last 150 years have been a period of relatively intense 
cattle grazing. There have been numerous homesteads in the valley, and agri- 
culture has been practiced on a limited scale on the Santa Cruz River flood- 
plain. The San Rafael Ranch was the location for the filming of many movie 
westerns, including the musical Oklahoma! 


METHODS 


Field work was conducted from August, 2000, through November 2001. Dur- 
ing the 17 months of the project a total of 62 trips were made, 34 to SCSNA and 
28 to SRSP. The field work thus covered three growing seasons: the summer/fall 
of 2000 and 2001 and the spring of 2001. The winter of 2000-2001 was unusu- 
ally wet, resulting in very productive collecting during spring of 2001. Since 
there was no direct vehicle access to the SCSNA, all of the collecting had to be 
done during long day-hikes. Plants were collected, mostly in duplicate, in plastic 
bags during the day and pressed either that same evening, or the following morn- 
ing after allowing collections to rehydrate somewhat. At SRSP most collections 
could be made on shorter hikes and pressed while still in good condition. 

A total of 1,286 specimens were collected, 689 from SCSNA and 597 from 
SRSP. Many trees, shrubs, and rosette succulents were not collected; rather their 
presence and distributions were recorded in field notes. Most cacti were identi- 
fied in the field using Benson (1969). A few species were included in the flora 
based on earlier collections from the study areas in the herbarium at the 


MCLAUGHLIN FIORA AREA AND SAN RAEAFI STATE PARK 669 


University of Arizona. Rick Gagnon, the ranger at SCSNA during the period of 
the field work, took hundreds of photographs of plants he encountered on that 
study site; a few species not encountered during the collection trips but photo- 
graphed by Gagnon are also included in the flora. 

In 2004, 1,064 ha adjacent to SCSNA, located to the northeast and upstream 
along the Coal Mine Creek watershed, was added to the park. This area, which 
is even more remote that the original acquisition, is likely to have additional 
taxa not found during the field work for this study. 

Geographic affinities of the floras of SCSNA and SRSP were examined by 
mapping the percentage of native taxa found in 245 local floras from Mexico, 
the United States, and Canada, using a database compiled by the author. Such 
maps use contours (isolines of percentage overlap) to show in a concise way 
what proportion of the flora extends into neighboring geographic regions. Three 
maps were prepared: one for native species occurring in the floras of both SCSNA 
and SRSP, one for those occurring only at SCSNA, and one for those occurring 
only at SRSP. 


RESULTS AND DISCUSSION 


Floristic Summary 

The total known flora for SCSNA is 561 species, 525 native and 36 exotic. For 
SRSP the total flora is 457 species, 407 native and 50 exotic. The percentage of 
exotic species in the flora of SCSNA is 6.4%, a typical value for floras in south- 
eastern Arizona, but low for local floras in most other regions of the United 
States. Exotics at SRSP constitute 10.9% of the total flora. There are 277 species 
in common to the two sites, 252 native and 25 exotic. The floristic similarily 
(Otsuka’s Index, based on native species only) between SCSNA and SRSP is 
54.5%, a somewhat low value for two areas in such close proximity (Fig. 1), indi- 
cating significant differences in the floras of semidesert and Plains grassland 
communities. 

There are 97 families and 354 genera in the flora of SCSNA, and 84 fami- 
lies and 274 genera at SRSP. The largest families in the two floras are listed in 
Table 2. The Asteraceae (composites), Poaceae (grasses), and Fabaceae (legumes) 
dominate in both floras, as they do in floras from most areas of the western 
United States. Even though the flora of SCSNA is larger than that of SRSP by 
more than 100 species, the number of Asteraceae are about the same and the 
number of Poaceae is actually larger at SRSP. The Poaceae also has the largest 
number of exotic species. The Euphorbiaceae is the fourth largest family at each 
site. One notable difference is the large number of Pteridaceae (ferns) at SCSNA, 
a family that is completely absent from the flora of SRSP Ferns in the Pteridaceae 
generally grow on rock outcrops or rocky slopes, a habitat not found at SRSP. 
The largest genera in the flora at SCSNA are Chamaesyce (Euphorbiaceae, 11 
species), Bouteloua (Poaceae, 8 species), Cyperus (Cyperaceae, 8 species), Dalea 


670 BRIT.ORG/SIDA 22(1) 


Taste 2. Largest families in the floras of Sonoita Creek State Natural Area and San Rafael State Park: 


numbers of genera (G), native species (NS), exotic species (ES), and total species (TS) 
Sonoita Creek State Natural Area San Rafael State Park 
Family G NS ES TS Family G NS ES TS 
Asteraceae 59 84 3 87 Asteraceae 46 79 3 82 
Poac 35 50 16 66 Poaceae 38 Di 18 75 
Fabaceae 31 51 2 53 Fabaceae 25 35 8 43 
Euphorbiaceae 11 230 (0 23 Euphorbiaceae 6 21 0 21 
Scrophulariaceae 11 15. 2 17 Brassicaceae 10 9 4 13 
Brassicaceae 13. 13—~«3 16 Solanaceae 6 1] 0 11 
Pteridaceae 7 13. (OO 13 Scrophulariaceae = 7 9 1 10 
Cactaceae 5 12 0 12 Cyperaceae 4 10 0 10 
Cyperaceae 4 12 O 12 Chenopodiaceae 4 7 1 8 
Convolvulaceae 3 11 0 11 Convolvulaceae 3 8 0 8 
Malvaceae 8 11 0 11 Malvaceae 5 / | 8 
Solanaceae 7 TO + 4 1 Onagraceae 3 8 0 3 


(Fabaceae, 8 species), Boerhavia (Nyctaginaceae, 7 species), Ipomoea 
(Convolvulaceae, 7 species), Brickellia (Asteraceae, 6 species), and Phacelia 
(Hydrophyllaceae, 6 species). At SRSP, the largest genera are Chamaesyce (9 
species), Bouteloua (9 species), Dalea (7 species), Ascle 7 (Asclepiadaceae, 5 
species), Euphorbia s.s. (Euphorbiaceae, 5 species), Ipomoea (5 species), 
Machaeranthera (Asteraceae, 5 species), and Oenothera ee 5 species). 

The percentage of the total flora (natives + exotics) that are epee: found 
only in aquatic and wetland habitats is 13.0% for SCSNA and 21.8% for SRSP. 
For those species found at SRSP but not at SCSNA the percentage of wetland 
plants is 34.8%, indicating the relatively greater importance of wetland habi- 
tats at SRSP. 


— 


Significant Taxa 

Ananthus squamulosa (A. Gray) King & H.E. Robinson (synonym Brickellia 
squamulosa A. Gray) (Asteraceae).—There is a relatively large population of this 
plant in Parker Canyon at SRSP. It is known in Arizona from only a few sites in 
the southeastern part of the state. 

Argyrochosma incana (C. Presl) Windham (Pteridaceae).—This is a rare fern 
in the United States, known only from a limited area in southern Arizona. It is 
uncommon on rock outcrops at SCSNA 

Cirsium grahamii A. Gray (Asteraceae).—This large, attractive, uncommon 
thistle is restricted to ciénega habitats. It occurs in the Heron Spring drainage 
on SRSP. 

Heterotheca rutteri (Rothrock) Shinners (Asteraceae).—This is a rare grass- 
land plant with a narrow range in southeastern Arizona. It is locally abundant 
at many localities in grasslands at SRSP. 


MCLAUGHLIN, FLORA OF REEK NATURAL AREA AND SAN RAFAEL STATE PARK 671 


Lilaeopsis schaffneriana (Schltdl.) Coulter & Rose ssp. recurva (A.W. Hill) 
Affolter (Apiaceae ).—This is a listed endangered species under the Endangered 
Species Act, the only such species so far documented from the floras of the two 
sites. It was previously recorded from Sharp Spring at SRSP. This species was 
not found at either Sharp or Heron springs, but a few populations along the 
Santa Cruz River were found at SRSP. 

Pennellia robinsonii Rollins (Brassicaceae ).—This species is new to the flora 
of Arizona and the United States. It differs from P. micrantha (A. Gray) Nieuwl., 
a woodland and forest species, in having shorter, wider siliques, dark purple 
sepals, and purplish-tinged petals (Rollins 1995). Rollins gives its range as re- 
stricted to the Mexican state of Chihuahua. The species is locally common in 
grasslands at SRSP. 

Phacelia sonoitensis S.P. McLaughlin ined. (Hydrophyllaceae).—A previously 
unknown Phacelia was collected several times at SCSNA. The plant occurs on 
rock outcrops or on talus below rock outcrops; it was found at several widely 
scattered sites. It hasa combination of characters that do not match any known 
Phacelia from the Southwest: pinnately compound leaves, narrowly spathu- 
late sepals, and a capsule of several shallowly cymbiform seeds. A description 
of this proposed new taxon has been submitted to Novon. 

Phyllanthus polygonoides Nutt. ex Spreng. (Euphorbiaceae).—This perennial 
herb is rare in Arizona, but is more common in Texas and Mexico. It is a rare 
plant at SCSNA in Fresno Canyon along the stream. 

Potentilla wheeleri S. Watson (Rosaceae).—This species occurs at SRSP; it is 
known from relatively few locations in southeastern Arizona, all at much higher 
elevations in adjacent mountains or from rock walls in steep-sided mountain 
canyons. Its occurrence at such a low elevation is remarkable. It was common 
on a north-facing slope adjacent to Sharp Spring. Several other high-elevation 
taxa occurred at the same site, including Thlaspi montana var. fendleri and 
Houstonia wrightii. 

Rhynchosia minima (L.) DC. (Fabaceae).—This is a new state record for Ari- 
zona. The species is common in northern Mexico and also occurs in the south- 
eastern United States. It is rare at SCSNA, found once on a rocky slope above 
Coal Mine Canyon and once in Fresno Canyon. 

Sibara virginica (Brassicaceae).—This is a new record for the state of Ari- 
zona; it was found on two occasions in dense sacaton stands adjacent to the 
Santa Cruz River in SRSP. Sibara virginica is a rather widespread species, and 
its status here as a native or exotic is not clear. 

Tephrosia vicioides Schlect.(Fabaceae).—This perennial herb was previously 
overlooked in the flora of Arizona, and is only known from the state from just a 
few collections. It was found in SCSNA ata single location on loose, rocky soil 
in the center of the par 

Tillandsia recurvata a) L. (Bromeliaceae).—The epiphytic species is com- 


672 BRIT.ORG/SIDA 22(1) 


mon in the tropics but is known in Arizona from just a few sites. It is locally 
common on junipers in narrow tributary canyons to Sonoita Creek at SCSNA. 


Floristic Geography 

Patterns of floristic affinity are illustrated by three maps (Fig. 4-6). These maps 
show the percentage of native species found in both parks (Fig. 4), found only 
at SCSNA (Fig. 5), and found only at SRSP (Fig. 6), which are present in each of 
245 other local floras from North America. The average range size (number of 
local floras out of the sample of 245 local floras) for native species occurring in 
both SCSNA and SRSP is 26.2 floras. These taxa extend throughout the Apachian 
floristic area (McLaughlin 1989, 1992) of southeastern Arizona, northwestern 
Sonora, southeastern New Mexico, and northwestern Chihuahua, with clear af- 
finities extending southeast into west Texas (Fig. 4). Ruderal species, wetland 
plants, and perennial grasses constitute a large proportion of the most wide- 
spread native species common to both parks. 

Those native species recorded only for SCSNA are more narrowly distrib- 
uted (mean range size = 15.6 floras) than those occurring in both parks. The 
average range size for the native species recorded only from SRSP is 25.2 local 
floras, similar to that for those species recorded for both floras. The higher 
proportion of wetland taxa among the native species found only at SRSP ac- 
counts in part for the greater average range size of these taxa. While the species 
found only at SCSNA are mostly Apachian in distribution, a relatively larger 
proportion extend into the Sonoran Floristic Province (McLaughlin 1992) of 
western Arizona, southeastern California, and throughout Sonora (Fig. 5). A 
much smaller percentage of those species recorded only from SRSP extend into 
the Sonoran Floristic Province, but a higher percentage extend southeast into 
the Madrean mixed-grass prairies (Bock & Bock 2000) of Chihuahua and 
Durango and east and northeast into the short-grass plains of eastern New 
Mexico and Colorado (Fig. 6). 

The climates of SCSNA and SRSP are not that different (Table 1); in both 
areas precipitation occurring during the summer monsoon (July to Septem- 
ber) accounts for 61% of the mean annual precipitation. The small differences 
in temperature and precipitation, however, are consistent with the relatively 
higher affinity of the warmer, drier SCSNA with the Sonoran Floristic Prov- 
ince, and for the higher affinity of the cooler, somewhat moister SRSP with the 
short-grass plains. The coarser textured, shallower soils of SCSNA may favor 
desert species while the deep clay soils of SRSP may favor species from the Great 
Plains. The floras of both parks, however, are overwhelmingly Apachian in their 


species com position 
I : 


MCLAUGHLIN, FLORA OF ONQOITA CREEK NATIIRA! AREA AND CAN RALCAEI STATE PARK 673 


55 
\ 2 
SO he, ean 
KO ees ‘a6 
A5- LP et 
id (i Gor i ‘ 
4 Lope 
40 aoe: 
Ce 
35- — a 
30 SAN i 
254 i 
. Catal Affinities of Speci 
ost 4 inities of Species 
o0- we ip. Occurringin both = L 
ay S, Sonoita Creek SNA 
<< and San Rafael SP 
15 T T T aoe ies 
-120 -110 -100 -90 -30 -70 
in both Sonoita Creek State Natural A 1 San Rafael S Park. Isolines 


Fic. 4. Affinities of nat 


based on a sample of 245 local floras. 


ANNOTATED CHECKLIST 


Nomenclature in the following checklist, with few exceptions, is based on 
Kartesz (1999). Nomenclature in Cactaceae follows Parfitt and Gibson (2003). 
Synonyms, when given, are for taxa listed under a different name in Arizona 
Flora (Kearney & Peebles 1960). Exotic species are preceded by an asterisk (*). 
Annotations for each species include: common name(s) when available, habit, 
collection numbers, and abundance and distribution for each study area (SC = 
Sonoita Creek State Natural Area; SR = San Rafael State Park). Spanish common 
names are those enco din northeastern Sonora, as listed by White (1948). 
Common names and habits are not repeated for varieties with the same data. 
Voucher specimens were deposited at the University of Arizona Herbarium 
(ARIZ); duplicates were used to establish field herbaria for the two state parks. 
Several species, including most cacti and succulents, many trees and shrubs, 
and species not found in flower, were identified in the field and not vouchered. 
Unless otherwise indicated, collection numbers are those of the author. 


BRIT.ORG/SIDA 22(1) 


Affinities of Species 


ccurring in 
_ Sonoita Creek SNA 
Only 
465 
I T T T T T 
-120 -110 -100 -30 -80 -70 
Fic. 5. Affinities of | 4 ry ly fa ra ft Lb C#eat iat LA ( z :< Cc Daf eae ees 
Park). Isolines represent the percentage of speci ly from SCSNA found in other local floras from North 
America, based on a sample of 245 local floras. 
LYCOPODIOPHYTA POLYPODIOPHYTA 

Selaginellaceae Azollaceae 
Selaginella rupincola Underwood. Spike moss; — Azolla mexicana C. Presl. Water fern; floating 

yerbaceous perennial.SC:8357; abundant on quatic. SC: 9330; rare in pools along Sonoita 


canyon slopes. 


EQUISETOPHTYA 


Equisetaceae 

Equisetum x ferrissii Clute (pro sp.).Scouring rush; 
herbaceous perennial. SR: 8268; asional, 
Santa Cruz River bottom 

Equisetum laevigatum A. Braun. Scouring rush; 
herb 8 


TA 
c 


eous perennial. SC: 8745; uncommon 
along Sonoita Creek. SR: 9432; common, 
Santa Cruz River bottom. 


Creek. SR: 8784, uncommon, Sharp Spring. 


Marsileaceae 

Marsilea mollis B.L. Robinson & Fernald. Pepper- 
wort; herbaceous perennial. SR: Niles 698 
(ARIZ); collected last in 1965 from a pool 
adjacent to the Santa Cruz River. 

Marsilea vestita Hook. & Grev. [M. mucronata A. 
Br]. Clover fern; herbaceous perennial. SC: 
8498, 8532, uncommon along streams In 
Fresno and Coal Mine Canyons. SR: Van 
Devender 84-443 (ARIZ);collected last in 1984 

from a pool adjacent to the Santa Cruz River. 


MCLAUGHLIN, 


AREA AND SAN RAFAEL STATE PARK 675 


T 


45-4 


40- 


35-4 


Affinities of Species 
Occurring in 
San Rafael SP Only 


18 T T 1 

-120 -110 -100 -90 -B0 -70 
Fic 6 ALE an 4 i + I a ul 1 L fa rs ic dR { £ j fa cr Le. Alas 1 
Area) | 1 al | ‘| £ f ly £ SRSP fi P| L 1 ims] L. L 
America kK ld | {7 cl im si fod ‘he Te 5 


Pteridaceae 
Adiantum capillus-veneris L. Maidenhair fern; her- 
ceous perennial. SC: 9419; below rock 

overhangs, west park boundary on south 
side of Sonoita Creek. 

Argyrochosma incana (K. Presl) Windham. Cloak 
ern; herbaceous perennial. SC:8407; uncom- 
mon on (i walls. 


n) Windham SSP. 
limitanea [Pellaea viva (Maxon) Mor- 
ton]. Cloak fern; herbaceous perennial. SC: 
8806; uncommon on rock walls. 

Astrolepis integerrima (Hook.) D.M. Benham & 
Windham [Notholaena sinuata (Lagasca 
Kaulf.var.integerrima Hook.]. Canahuala; her- 
baceous perennial. SC: 8360, 8454; uncom- 
mon on rock walls. 


a 


Astrolepis sinuata (Lagasca ex Swartz) D.M. 
R Kk PO \AL 1} M TAT L | 


ra 


sinuata (Lagasca) Kaulf. var. sinuata]. 
sae herbaceous perennial. SC: 8406; 
mmon on rocky slopes and rock walls. 

Sees ia hispida (Mett. ex Kuhn) Underwood. 
A ommeria; herbaceous perennial. SC: 
8400; uncommon, mostly below oaks. 

Cheil sna eatonii Baker. Lip fern; ieee 
perennial. SC: 9677; rare, canyon walls. 

Cheilanthes lindh k. Fairy swords; her- 
baceous perennial. SC: 8380; common on 

slopes and rock walls. 

Chei ne wrightii Hook. Wright's lip fern; her- 
baceous perennial. SC:8416,8820,8871;com- 
mon on rocky slopes 

Notholaena grayi Davenport. Cloak fern; herba- 


elmer) Hoo 


ceous perennial. SC: 8357; uncommon on 
rock walls 

Notholaena standleyi Maxon. Cloak fern; herba- 

eous perennial. SC: 8410, 8822; uncommon 
on rock walls. 

Pellaea intermedia Mett.ex Kuhn. Cliff brake; her- 
baceous perennial. SC: 6846, 9217; rare on 
ny is 

Pellaea wrightiana Hook. Cliff brake; herbaceous 
perennial. SC: 8878, 8933, 9217, rare on rock 

walls. 


PINOPHYTA 


Cupressaceae 
| , L 


‘ 7 NA 7 pp 


Adams (J. monosperma (Torr.) Little sensu 

Kearney and Peebles (1960), i 

huata, tascate, tdscale; small tree. SC: 9408; 
nin canyons, uplands 

Juniperus deppeana Steud. Alligator juniper, 

huata, tascale, tascate; small tree. SR: 9392; 

uncommon,grasslands east of Santa Cruz River. 


n part]. Juniper, 


MAGNOLIOPHYTA: MAGNOLIOPSIDA 


Acanthaceae 
Anisacanthus thurberi (Torr.) A.Gray. Desert hon- 
re large shrub. SC: 9280; commen in 
anyon 


cago arizonica A. Gray. Herbaceous pe- 
nial. SC: 9291, 9297; uncommon in mes- 
quite gra ae ds. 


Sia resupinata (Vahl) Juss. Herbaceous 
f 


nial. SC: 8734; common in shade o 
gallery forests along Sonoita Creek. 
ees decumbens (A. Gray) Kuntze. Herba- 
ceous perennial. SR: 8287; common, grass- 
lands. 
Elytraria imbricata (Vahl) Pers. P Purple scaly stem, 
cordoncillo; herbaceous perennial. SC: 8369; 


anyons, rocky slopes. 
aenetate ium nervosum Nees [7. hispidum Nees]. 
Herbaceous perennial. SC: 8348, 9232; un- 
common, mostly in canyon bottoms. 


Aizoaceae 
Trianthema portulacastrum L. Horse pveRne 
summer annual. SC: 841 9;0c over- 


grazed sites. SR: 8791; uncommon, sacaton 
stands 


BRIT.ORG/SIDA 22(1) 


Amaranthaceae 

Alternanthera pungens Kunth [A. repens (L.) 
Kuntze]. Khakiweed; mat-forming herba- 
ceous perennial. SR: 87/2; occasional, grass- 
lands. 

Amaranthus palmeri S.Watson. te aay 
bledo, quelite; summer annual. 3c: 8352 
common, canyons at 
8270, common, mostly along drainages. 

Amaranthus torreyi (A.Gray) Benth. ex S.Watson 
Torr.’s amaranth; summer annual. SC: 8392, 
8422, uncommon, mostly in canyons. 

Froelichia arizonica Tho 
cotton; herbaceous ete SC: 8809; rare, 
found once in Fresno 

COMP aiena caespitosa i Globe almakenisiy 


* a 1] Cc | 
rnbe rex standley. snake 


CD. O072 


grasslands:S E 
Gomphrena nitida Rothrock. Globe amaranth; 
herbaceous perennial. SC: 8487; common, 
10; occasional, grass- 


grasslands 


grasslands. SR: 8283, 83 
lands. 
Gomphrena sonorae Torr. Globe amaranth; her- 
aceous perennial. SC:8370; common, grass- 


ei OF 


nds. 
Guilleminea densa (Humb. & Bonpl. ex Willd.) 
Mog. [Brayulinea densa (Humb. & Bonpl.) 


Small]. H 
ianal 


baceous perennial SC: 8415; 0cca- 
J Cr. Sap of | pee 


Iresine heterophylla Standley. Bloodleaf; herba- 
C ; 


eous perennial. SC: 8751; rare, canyons 


Anacardiaceae 
Rhus trilobata Nutt. var. anisophylla (Greene) 
Jepson. Sumac 


9630; rare, oak woodland adjacent to Parker 


quawbush, lima; shrub. S 


anyon. 

Rhus trilobata Nutt. var. pilosissima Engelm. 
Shrub. SC: 9209; uncommon, canyons 

Rhus virens Lindheimer ex A. Gray var. 
choriophylla (Wooton & Standley) L.Benson 
[R. choriophylla Wooton & Standley]. Lima; 

rare, canyons. 

Toxicodendron rydbergii (Small ex Rydberg) 
Greene [Rhus radicans L.var.rydbergii (Small) 
Rehder]. Poison ivy, hiedra; woody vine. SC: 
9420; uncommon, canyons. 


Apiaceae 
Berula erecta (Huds.) Coville. Water parsnip; her- 
baceous perennial. SR:8276;commMon, springs. 


MCLAUGHLIN, FLORA OF 


AREA ANN CAN RAEAFI STATE PARK 


Bowlesia incana Ruiz & Pavoén. Hairy bowlesia; 
winter annual. SC:8875;abundant, especially 
below mesquites. 

*Conium maculatum L.Poison hemlock; biennial 

C: 9474; rare, along Sonoita Creek. 

Daucus pusillus Michx. American carrot; winter 

annual. SC:8926;common on rocky uplands. 


ro 


herbaceous perennial. SR: 8375; occasional, 

grasslands. 

Hydrocotyle ranunculoides L.f.Water pennywort; 

atic herbaceous perennial. SC:8496; rare, 

pools in Fresno and Coal Mine Canyons. SR: 
8787, occasional; Santa Cruz River bottom. 

Lilaeopsis schaffneriana (Schitdl.) J.M. Coulter & 
Rose var. recurva (A.W. Hill) Affolter. Huachuca 
water ae inconspicuous parce 
peren SR: 9434 allo 
ter fe sont Cruz River. 

Lomatium nevadense (S.Watson) J. M.Coulter & 
Rose var.parishii J.M.Coulter & Rose) Jepson. 
Indian root; herbaceous perennial. SC: 8843, 


9084; rare, mesquite grasslands 
Spermolepis echinata (Nutt. ex DC.) Heller. Scale 

seed; winter annual. SC:8903; occasional, can- 

yons and uplands. SR: 91704; uncommon, 


grasslands. 

Yabea microcarpa (Hook. & Arn.) K.-Pol [Caucalis 
microcarpa Hook. & Arn.].Winter annual. SC: 
9024, 9025; locally abundant, north-facing 
slopes in oak savanna. 


Apocynaceae 

Apocynum cannabinum L. Dogbane, Indian 
hemp; herbaceous perennial. SC: 9360; rare, 
one population in Coal Mine Canyon 

Haplophyton crooksii (L.Benson) L.Benson.Cock- 
roach plant, hierba de la cucuracha; herba- 
ceous perennial. SC: 9500, E.A. Lewis 012; rare, 

slopes, canyons 

Telosiphonia brachysiphon (Torr.) Henrickson 
ee brachysiphon (Torr.) A.Gray]. 
Rock trumpet; herbaceous perennial. SC: 
8523, a Lewis 013, locally abundant on 
rocky slopes. SR: 9579; rare, grasslands 


Araliaceae 
Aralia humilis Cav. Tepetate; shrub. SC: 9703; rare, 
cky north-facing slope 


Aristolochiaceae — 


WW, & Standley. Indian 


Ar istoloct HCA VV CACIUTETT 


root; herbaceous perennial. SC: 8907, 9550; 
uncommon and infrequent, mostly on rock 
walls. 


Asclepiadaceae 

Asclepias asperula (Dcne.) Woodson ssp.asperula 
(A. capricornu Woodson sensu Kearny and 
Peebles (1960)].Antelope horns; herbaceous 
perennial. SC: 9220; rare, grasslands. SR: 9397; 


uncommon, grasslands 
lepias involucrata oe Milkweed; herba- 
ceous perennial.SR:9154 gras 
lands. 


apace inaria Cav. Milkweed; shrub. SC: 6437; 
rare, r 

Asclepias ae a Torr. Milkweed, yerba de 
cuerve; herbaceous perennial. SC: 9279; rare, 
grasslands, rocky slopes. SR: 9394; occasional, 
grasslands. 

Asclepias nyctaginifolia A.Gray.Four o'clock milk- 
weed; herbaceous perennial. SC: 9551; rare, 
canyons, rocky slopes. SR: uncommen, grass- 
lands. 

Asclepias subverticillata (A.Gray) Vail. Poison milk- 

weed, orled milkweed; herba- 
eOus sect SR: 8239; common, grass- 
onde Sharp Spring drainage. 

Asclepias tuberosa L. ssp. interior Woodson. But- 
terfly weed, pleurisy root, orange m |kweed; 


grasslands. 

Funastrum cynanchoides Decne. ssp. hetero- 
payin (Engelm. ex Torr.) Kartesz [F 
het wllum (Engelm.) Standley].Climbing 
siloeee herbaceous Peleg, vine. SC: 
929): rare, cCanyo 
shrubs in sec 


,on 


Asteraceae 


A.Gray) Reveal & King [Perezia 
compe A. Gia Large herbaceous peren- 
Se 8771; rare, canyons 
C Gray) fave & King [Perezia 
wrightii A.Gray]. Brownfoot; herbaceous pe- 
rennial. SC: 8922; rare, uplands 
Ambrosia confertiflora DC. [Franseria confertiflora 
C.) Rydberg] SU ea ues, herbaceous 
perennial. SR: 8 grasslan 
Ambrosia pallostachy DC. Western cel 
h us perennial.SR:8599; locally abun- 
dant ae Parker Canyon. 


ACOuUI CU tl er beri 


= 


OWI wrig! PUT i (A. 


678 


Ambrosia trifida L. var. texana Scheele [A. oe 
DC.].Ragweed, chuchuni; herbaceous peren 
nial. SC:8656;abundant along Sonoita Creek. 
SR:8302;common, roadsides, low areas, Santa 
Cruz River bottom 

Ananthus squamulosa (A. Gray) King & H.E. Rob- 
inson [Brickellia squamulosa A. Gray]. Herba- 
ceous perennial. SR: 9728, 9736; locally com- 
mon, Parker Canyon. 

Antheropeas lanosum (A. Gray) Rydberg. 
[Eriophyllum lanosum A. Gray].Woolly-daisy; 
winter annual. SC: 8920A, 9076; common in 
mesquite grasslands. 

Artemisia campestris L. ssp. borealis (Pallas) Hall 

lements var. scouleriana (Hook.) Cronq.[A. 
pacifica Nutt.]. Wormwood; herbaceous pe- 
rennial. SR: 9623; rare, Parker Canyon 

Artemisia dracunculus L. [A. dracunculoides 
Pursh]. False tarragon; herbaceous perennial. 
SC:970/, rare, north-facing slope in Coal Mine 


Canyon. 
Artemisia ludoviciana Nutt.ssp.a/bula (Wooton) 
Keck \A/ 1.1 | pee SC: 


8673; common, north-facing slopes. SR: oc- 
casional, oak woodlands and savannas. 
Artemisia ee Nutt. “SSP. eae (Rydberg) 

Keck _oak woodlands 


and savannas. 


ii A. Gray. Shrub. SR: 9694, 9726; 
rare,Oa siAwoudiand adjacent to Parker Canyon. 

ee DC. Yerba de pasmo; small 
shrub ommon in grasslands. SR: 
9376; asslands 

Baccharis eae ia ete & Pavon) Pers. [B. 
glutinosa Pers.].Seep willow, batamote jarilla; 
shrub. SC: 9651; occasional along Sonoita 
Creek, Fresno Canyon. SR: 9660; occasional, 
Parker C n. 

Baccharis ere A, ey Desert broom, 
romerillo; shrub. SC:881 1 

common, mostly roadsides. 

ee eines Kunth. Shrub. SC: 8759; rare, 

canyon one! north-facing slopes. 
ANIA 


> 


common, canyons 


absinthifolia Benth. var. dealbata (A. Gray) 
A. Gray. Perennial herb. SC: 9277; rare, rocky 
slopes. 


Baileya multiradiata Harvey & A. Gray ex A. Gray. 
Desert marigold; perennial herb. SC: 9742; 
rare, grasslands. 

Barkleyanthus salicifolius (Kunth) H.E. Robinson 


BRIT.ORG/SIDA 22(1) 


& Brett. [Senecio salignus DC] Shrub.$C:8974; 
common in gallery forest along Sonoita 


reek, 

Bebbia juncea (Benth.) Greene sss cnuseeds 
Sweet bush;shrub.SC:9414:ra 

Berlandiera lyrata Benth. Cheese flower ae 
de la vibora; herbaceous ie SR: 9160; 
occasional, grasslands, roadside 

Bidens aurea (Ait.) Sherff. Bur eee peren- 
nial herb. SC: ‘as 8773, 9719; uncommon, 
along stream 

Bidens bigelovil a Gray. Beggar ticks, Spanish 
edles; summer annual. SR: 9619, 9692; un- 
n, Parker Canyon. 
nape ferulaetot Waed) DC sbegdal WeNs apa: 


Bidens laevis (L.) B.S.P. Bur marigold; herbaceous 
pere ae SR: 878 | 73; common, springs, 
z River bot 
Pe ena met on seal ticks, Spanish 
needles; summer annual. SC:8356;abundant, 
canyons, oak woodlands. SR; 8700, common 
below oaks. 
Brickellia amplexicaulis B. L. Robinson var. 
amplexicaulis. Perennial herb. SC: 8763; rare, 


upper Fresno Canyon. 
| lia baccharidea A. Gray. Shrub. SC: 9680; rare, 


o Can 
een. nena ia A.Gray. Herbaceous peren- 
nial. SR: 9699; rare, oak sav 
Brickellia californica (Torr. & < Ges) A. Gray vat. 
californica. Shrub. SC: 8807; uncommon, 


canyons. 
Brickellia coulteri A.Gray. Perennial herb. SC:8837; 
rare, canyon 
Brickellia eupat are (L.) Shinners. var. 
chlorolepis (Wooton & Standley) B.L. Turner 
[Kuhnia rosmarinifolia Vent. var. chlorolepis 
(Wooton & Standley) Blake]. Herbaceous 
perennial. SR: 8563; common, grasslands. 
Brickellia floribunda A. Gray. Robust perennial 
herb. SC: uncommon, along Sonoita Creek. 
SR: 9717; common, Parker Canyon 
Brickellia venosa (Wooton & Standle B. L. Rob- 
n.Herbaceous perennial.SC:8676B, 9720; 


Me eee ons 


Calycoseris wrightii A. Gray. Tackstem; winter an- 
nual. SC: 9074; rare, uplands. 

DC. Plume weed; summer 

annual. SC: 8657; rare, shaded canyon walls. 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL 


AREA AND SAN RAFAEL STATE PARK 679 


Carphochaete bigelovii A. Gray. Bristlehead; low 
shrub. SC: 9283: rare, rocky uplands. SR: 9102; 
rare,oak woodland adjacent to Parker Canyon. 

Chaetopappa ericoides (Torr.) Nesom [Aster 
arenosus (Heller) Blake, A. hirtifolius Blake]. 
White aster, rose heath; herbaceous peren- 
nial. SC: 9029; occasional, mesquite grass- 
lands. SR: 9377; grasslands 

Cirsium gana A. Gray. Thistle; large biennial. 

uncommon, springs. 

Cirsium neomexicanum A. Gray. Thistle; biennial. 

: 9136; occasional, mostly in canyons. SR: 
oscasionial eee 


Vall ; (Bees | 


biennial. SR: 9368; common, aeacs| ands. 

Conyza canadensis (L.) Cronq. [Erigeron cress 
L.]. Horseweed; summer annual. SC: 8 
common, canyons. SR: 8696; locally ae 
dant, grasslands. 

Cosmos Brees (Jacq.) Pers.Cosmos;summer 
annual. SR: 9714; rare, Parker Can 

Eclipta eal. .) L.[E. alba (L.) Tee Yerba 
tajo; summer annual. SC: 9669; rare, wet 

along Sonoita Creek. 

Ericameria taricifolia (A. Gray) Shinners 
[Aplopappus laricifolius A. Gray]. Turpentine 
bush; shrub. SC: 9730; common, north-facing 
slopes Caen 

Ericameria nauseosa 5 Pals ex Pursh) ow & 

aird ssp.nauseosa var.latisquamea A.Gray) 

Nesom & Baird en a nauseosus 

(Pallas ex Pursh) Britt. ssp. fatisquameus (A. 

ay): Hie & einen) Rabbitbrush; shrub. 
P n 


4: abur Wd tlt, 


a8 


Sieeron ariosolus Nesom [E. divergens Torr. & A. 
Gray, in part]. Fleabane; annual or herba- 
ceous perennial. SR: 8237, 8279, 8549B; com- 
mon, grasslands. 

Frigeron colomexicanus A. Nelson [E. divergens 
Torr.& A. Gray, in Ae Fleabane; herbaceous 

ncommon, along stream, 


perennial.SC: 9271 


resno Can 

eerie ten Torr.& A.Gray. Spreading flea- 
bane; annual or herbaceous ees SE 
he oe 908?,9224;com yons and 
uplands.SR: 9700, roe concn toa 

Erigeron flagellaris A.Gray.Fleabane; herbaceous 
perennial. SR: 9246, 9252, common, 
grasslands and sacaton flat 

Erigeron oreophilus Greenm. Ftabere herba- 


ceous perennial. SC: 8477, 
yons, below 

Evax verna Raf. var. verna [E. multicaulis DC. var. 
multicaulis]. Rabbit tobacco, cotton rose; 
winter annual. SC: 9073; common, mesquite 
grasslands. SR: 9710, 9250; occasional, grass- 
ianiGS 

Fil ica Nutt. Yert i annual. 


SC: 9014B, 9017, 9087;common, canyons and 


_can- 


slopes. 

Fleischmannia pycnocephala (Less.) King & H.E. 
ROpInSen LEP acorn pycnccenagiom 
Less ].T 


SC: 8774; rare, north: facing slo opes. 

Gaillardia pinnatifida Torr. Blanket flower; herba- 
ceous perennial. SR: 8726; occasional, grass- 
lands and roadsides 

Gamochaeta purpurea (L.) Cabrera [Gnaphalium 
purpureum L.]. Cudweed; winter annual. SC: 
. ci 9026; abundant, mesquite grasslands. 

“common, grasslands. 

eneee eee A. Gray. Shrub. SC: 9552; 
uncommon, Fresno Canyon. 

ag alee eee (DC.) A. Gray. 

nakeweed; subshrub. SC: 8772; common in 
ee ; 

Helenium thurberi A. Gray. Sneezeweed; annual. 

SC: 8659, 8747; uncommon along streams. 

i th | (é f] ; | 


J>/ 


tall summer annual. SC: 9653; occasional, 
along Sonoita Creek. SR:8303; abundant, low 
areas, roadsides. 

Helianthus petiolaris Nutt. Sunflower, girasol; 
summer annual. 3; rare, canyons. SR: 
8731;common, Parker Can 

ayaa longifol ia eo. & Greenm.) 
Cockerell var. annua (M.E. Jones) Yates 
[Viguiera annua (Jones) Blake]. ean gold- 
eneye; late summer Falpual ae eke com- 
mon, uplands 

Heliomeris multiflora Nutt. ee “multiflora 
(Nutt.) Blake]. Golden eye; herbaceous pe- 
ee 

Heterosperma pinnatum Cav.Summer annual. SC: 

common, north-facing slope, Coal 
Mine Canyon. SR; 8590, common, Parker 


anyon. 
Heterotheca rutteri (Rothrock) Shinners. Herba- 
erennial. SR: 8264; locally common, 

grasslands. 


680 


Heterotheca subaxillaris 
plant; herbaceous perennial. SC: 
ae common, along Sonoita Creek, can- 
yons. SR: 8553; common, grasslands and 
roadsides. 
EOSES mencgyd Torr. & A. Gray ex A. Gray. 
8812; common along 
Sonoita Creek. SR: 9693; rare, Parker Canyon. 
Ba wislizenil A. Gray. Summer annual. 
583; occasional, grasslan 
pienoninwe ightli A.Gray. cr ean 
nial. SC: 8768, 9732; uncommon, Fres 


a 


| | 
Bul robush; PIRI Bele SC: 


lsocoma tenulsecta Greene [Aplopappus 
tenuisectus potas Blake]. Burroweed; sub- 
shrub. 9707, common, ers aed 
nds. SR: 971 3; uncommon, gq 
etal lon solidaginifolium (A. se ca & 
Robin Sa eel iallal, daginifol ow 


wih 


s (Lam.) Britton & Rusby. 


BRIT.ORG/SIDA 22(1) 


Machaeranthera tanacetifolia (Kunth) Nees 
ter tanacetifolius (Kunth) Nees]. oe 
perennial. SC: ; uncommon, along 
ie Creek. SR: 8343, 8703, pe 9677. 
common, roadsides, Parker Canyon, Santa 
Cruz floodplain 

Malacothrix clevel andl A. Gray. eae casa 
saucers; winter annual. SC: 9079, 9121; un 
common, canyons and bees Jones SR: 9242; 
uncommon, mostly below 

Malacothrix fendleri A. fe or dandelion; 
winter annual. SC: 9015; common, me i 
arasslands. SR:9097 9157 clans 


MmMon 


Melampodium longicorne A. Gray. Summer an- 
nual.SC:8385; uncommon, canyons. SR:85917; 
tale Pane! Ca YOR. 


g.[M hispid } h]. 

Summer aes oe 8663; uncommon, can- 
yons. SR:8309, 8701: arker Canyon. 
Packera (A. ani [ A.Weber & A. 


Gr 
SC: 9506, 9731: ;rare, north-facing slopes. 
*“Lactuca serriola L. Prickly lettuce, wild lettuce; 
nnual. ee, eee Saas rocky 
slopes. SR nds 
Laennecia euler (A. Gray) Nesom [Conyza 
coulteri A. = SUING annual. SC: 9649; 
uncommo along Sonoita Creek. 
SR: 8338; oc fasta 
Laennecia sophiifolia (Kunth) Nesom sai ied 
sophiaefolia Kunth]. Annual. SC: 8765 
common, canyons. SR: 9657; eal 


gr 


grasslands. 

Lasianthaea podocephala (A. Gray) K. Becker 
[Zexmenia podocephala A. Gray]. Herbaceous 
perennial. SC: 85 
canyons. SR: 9629; rare, oak woodland adja- 
cent to Parker Canyon. 

Machaeranthera gracilis (Nutt.) Shinners 
[Aplopappus gracilis (Nutt.) A. Gray]. Annual. 
ep G009, common, ioe uplands. SR: 


5 9583:rare, below trees in 


filachaerantherani aril eet ) Shinners ssp 
innatifida var. pinnatifida [Aplopappus 
spinulosus (Pursh) DC. var. turbinellus 
(Rydberg) Blake]. abnectars ems SR: 
paul gO common, grass 


[Act 


agetinus 
(Greene) Blake]. Hereaeeous. pee SC: 
8395, 9586; common, below mesquite. SR: 
9572: al grasslands 


Love Gace neomexicanus A. Gray]. Hew 
NA SR: 


9387. rare, drainages. 
Parthenice mollis A.Gray.Robust summer annual. 
rare, canyons 
Pectis ie Harvey & A, Giay Fetid marigold; 
summer annual.SC:8477; uncommon, uplands. 
ae longipes A. Gray. Fetid marigold; herba- 
ceous perennial. SC: 9228; uncommon, mes- 
quite grassland. SR: 8777, 9375; uncommon, 
grasslands 
Pectis prostrata Cav. Fetid marigold;summer an- 
ual. SC: 8537; uncommon, grasslands. SR: 
9618; rare, Parker Canyon 
ec gracile Benth. Galen: herbaceous 
perennial. SC: 9364; rare, uplan 
ee um ruderale (Jacq.) a SSp. 
macrocephalum i. R.R. Johnson [P 
san eee um DC]. Yerba del venado;sum 
mer annual. SC: 8423; uncommon, canyons. 
Se aie canescens (DC.) W. A. Weber 
ssp.canescens [Gnaphalium wrightii A. Gray]. 
Cudweed; eae perennial. SC: 8766; 
uncommon, canyons. SR: 8318, 8699: locally 
mon, grasslands 
eee di leucocephalum (A. Gray) A. 
Anderb. [Gnaphalium leucocephalum A. 
Gray]. White cudweed; herbaceous peren- 
nial. SC:8388, 8755; uncommon, canyon bot- 
tom. SR: 9697; uncommon, Parker Canyon. 


=a 


MCLAUGHLIN, FLORA OF 


RAL AREA AND SAN RAFAEL STATE PARK 681 


Pseudognaphalium pringlei (A. Gray) A. Anderb. 
as ium pringlei A.Gray].Cudweed;her- 
baceous perennial. SR: 9519; occasional, 

a 
Pseudognaphalium stramineum (Kunth) W. A. 
r [Gnaphalium chilense es Cud- 
weed: annual.SC: 9 an 
SR:8267; occasional, Santa Cruz River es 
aa ate aes A. Gray. Summer annual. SR: 
t,low areas, roadsides, washes. 

Buinesanees pauciflorus (D. Don ee 
multicaulis DC.].False dandelion; hereacdeus 
perennial. SR: 8305 


or 


8 Q2 
2} i UNICO TIOTT, 


rings and Santa Cruz River bottom 
ica: a californica Nutt. California saicony 
winter annual. SC: 9362; uncommon, rocky 


slopes. 

Rafinesquia neomexicana A.Gray. Desert chicory; 
winter annual. SC: 9012, 9016; common, can- 
yons, rocky slopes. 

Sanvitalia abertii A. Gray. Summer annual. SC: 
see rare, canyons. SR: 8778, 9625; uncom- 

on, Parker Canyon 

oe iO oe idus oe val. enaceaus [S.longilobus 
Ben f groundsel;shrub. SC: 9284; 

les canyons. SR: 8778; occasional, 
grasslands. 
olidago velutina DC.[S. sparsiflora A.Gray].Gold- 
enrod; herbaceous perennial. SC: 9/12; rare, 
one patch found in canyon west of Fresno 
Canyon. SR: 8578; uncommon, mostly oak 
woodlands 

*Sonchus asper (L.) Hill. Sow thistle; annual. SC: 
9151; occasional, mostly in canyons. SR: 9156, 

1 joccasional Santa Cruz River bottom. 

Nit lA latt 4 | 


ce 9471: rare, canyon 
sepharomea pcr ci A. Bd He 
erennial. S Oc- 


lettuce; herbac 
casional, canyons 7 ce ms os ae 

occasional, grasslands 
ee thurberi A. Gray. Wire lettuce; 
mer annual. SR: 9450; occasional, grass- 


an 
ee oan divaricatum . ee Nesom [As- 
ter exilis Ell.]. Annual. SC: 8 common in 
ere ene LM 
River bottom. 
Symphyotrichum falcatum (Lindley) Nesom var. 
commutatum (Torr. & A. Gray) Nesom [Aster 


commutatus (Torr. & A. Gray) A. Gray]. White 
prairie daisy; perennial herb. SR: 8577; com- 


mon, springs. 

*Taraxacum officinale G. H. Weber ex F.H.Wigg. 
Dandelion; annual. SC: 9475; rare in wet 
ground along streams. SR:97712;uncommon, 
prings. 


at enn megapotamicum (Spreng.) Kuntze. 
Her ous perennial. SR: 9369; common, 
aie a 
Thymophylla pentachaeta (DC.) Small var. 
hartwegii (A. Gray) Strother [Dyssodia 
pentachaeta (DC.) B. L. Robinson]. Herba- 
eOUS perennial SC: 9271,9?72.uncommon, 
uplands. 
tea ia POE A. Gray. Summer annual. SC: 
north-facing slopes. 
Towinceneis (Richardson) Porter. Herba- 
Cceous sefeitial SR: 9257; rare, grasslands 
Trixis californica Kellogg. shrub. SC: 8364; occa- 
sional, canyons and rocky slop 
Tee: lindleyi (DC.) ee Gea: 
linearifolia (DC.) Schultz-Bip.]. Silver puffs; 
winter annual. SC: 8867, 8924; occasional, 
uplands. SR: 9054 |, grasslands, be- 


low oaks. 
Koc hioid, 


v.) Benth. & Hook.f. ssp. 
exauriculata (Robinson & Greenm.) J.R.Cole- 
Crownbeard;summer annual. SC:86917; 
weedy terrace adjacent to 

Sonoita Creek. SR:uncommon, Parker Canyon. 
Viguiera cordifolia A. Gray. Golden eye; herba- 
eous perennial. SR: 8598; locally common, 


Viguiera dentata (Cav.) Spreng. var. dentata. 
Golden eye; Ded perennial. SC: 8672; 


abundant in mesquite bosques adjacent to 
Sonoita Creek. SR: one sbunaant oak 
odlands 


OQI4 
. 00. 


Viguiera dentata (Cav.) Spreng. var. lancifolia 
i | ro 


Xanthium strumarium L. var. canadense (P. Mill.) 
Torr. & A. Gray [X. saccharatum Wallr]. Cock- 
lebur,abrojo;summer annual. SC:8758; abun- 
dant in moist ground along streams. 5R:8597; 
common, low areas, washes, Santa Cruz River 


ottom. 
Xanthocephalum gymnospermoides (A. Gray) 
th. & Hook. Summer annual. SR: 8262; 


abundant, grasslands 


pak 


Zinnia acerosa (DC,) A. Gray [Z pumila A. Gray]. 
ert zinnia; low shrub. SC: 9326; uncom- 


mon, uplands. 
Zinnia grandiflora Nutt. Prairie zinnia; herbaceous 
erennial. SR: 9400 ional | 


grasslands 


bas 


Zinnia peruviana (L.) L. [Z.multiflora LJ]. Summer 
annual. SR:8720; uncommon, Parker Canyon. 
Berberidaceae 
Berberis wilcoxiik Bart “shrub. SC: 9276; 
rare, north-facing slope in Fresno Canyon. 


Bignoniaceae 

Chilopsis linearis (Cav.) Sweet ssp. arcuata 
(Fosberg) Henrickson. Desert willow, mimbre; 
small tree. SC: 9468; uncommon, along 
Sonoita Creek. SR: 9621; abundant, Parker 
Canyon. 

Tecoma stans (L.) Juss.ex Kunth var.angustatum 
Rehder. Trumpet flower, l/uvia de oro, palo de 
arco, gloria; shrub. SC:882]:rare, rocky slopes. 


Bixaceae 
Amoreuxia palmatifida Moc. & Sessé ex DC. 
lic ue perennial herb.£.A. Lewis 071; 
mon, grasslands and uplands. SR: 9526; 
ae cee 


Boraginaceae 

Amsinckia menziesii (Lenm.) A. Nelson & J. F. 
Macbride var. intermedia (Fischer & C. A. 
Meyer) Ganders IA. intermedia Fischer & C.A. 
Meyer]. Fiddleneck; winter annual. SC: 9021; 
rare, Coal Mine Canyon. 

Cryptantha angustifolia (Torr) Greene. Narrow- 

annual. SC: 


i barbigera (A. Gray) Greene. Bearded 
popco ower; mainte anvil SC:8902, 8906 
canyons and uplands. 


8925, 9 
SR: aie rare, oak savannas 

so ceerate micrantha (Torr) |. M. Johnston var. 
micrantha. Purple-rooted popcorn flower: 
ence SC: 9133; ndy 
floodplain of Sonoita Creek. 

Cryptantha pterocarya (Torr.) Greene var. 
cycloptera (Greene) J. F. Macbride. Winter 
annual. SC: 8866, 9179: abundant, mostly on 

ocky slopes. 

Cryptantha pusilla (Torr. & A. Gray) alg Pop- 
corn flower; winter annual. SC: 8865, 9069, 
9139B; uncommon, canyons and sl up- 
lands. SR: 9755; uncommon, grasslands. 


BRIT.ORG/SIDA 22(1) 


Lithospermum cabrense Greene. Puccoon; herba- 
ceous perennial. SR: 9482, 9486; common, 
grasslands 

Lithospermum incisum Lehm. Puccoon; herba- 

eous perennial. SR: 9302; occasional, grass- 
lands. 

Pectocarya recurvata |.M. Johnston. Arch-nutted 

bbur; winter annual. oe aed wel 9132; oC. 


casional, canyons, slopes. S 


grasslands, disturbed areas. 
Plagiobothrys arizonicus (A. Gray) Greene ex A. 
Gray. Blood weed, Arizona popcorn a 
winter annual.SC:8855,9116;common,gra 
lands and canyons. SR: 894 
ton flats, grasslands. 


/$aca- 


Brassicaceae 

Arabis perennans S. Watson. Rock cress; herba- 
ceous perennial. SC:8831,8858; uncommon, 

walls. 

eo bursa- patel (L 
purse, 
rare, gallery forest along Sonoita sae . 
9047. disturbed areas. 

Descurainia pin (Walt.) Britton ssp. 
‘an a Detling. Tansy mus- 
tard; winter annual. SC: ;common, can- 
yons and uplands. SR: 9058; abundant, road- 
sides, grasslands 

Descurainia pinnata (Walt.) Britton. ssp. 
ochroleuca (Wooton) Detling. Tansy mustard; 
winter annual. SR:8939; abundant, roadsides, 
grasslands. 

“Descurainia sophia (L.) Webb ex Prantl. Flixweed; 
winter annual. SC: 9137; uncommon, along 

9048 oadsides 


.) Medik. oe S 


panique illo; winter annual. S 


asslands. 
Draba cuneifolia Nutt. ex Torr. & A. Gray var. 
a. S.Watson. Whitlow grass; winter 
nnual.SC: 8826, 8840; common, canyons. SR: 
ae uncommon, clara 
Gray. Winter annual. 
Ne 88/73, 9013; rare, oe walls, Fresno and 
Coal Mine canyons. 
*Lepidium campestre (L.) Ait. f. Peppergrass; win- 
ter annual. SR: 9300; rare, springs 
Lepidium lasiocarpum Nutt. var. lasiocarpum. 
Sand peppergrass; winter annual. SC: ioe 
commo land d drai SR:8 


Dry opeta lo inathim 


abundant, ‘low areas, grasslan 
Lepidium ie Wooton. Spalciny spring- 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL 


AREA AND SAN RAFAEL STATE PARK 683 


summer annual. SC: 9491; uncommon, can- 
yons. SR: 9303; locally abundant, Parker Can- 
yon, grasslan nds. 

Lepidium virginicum L.var.medium (Greene) C.L. 
Hitchcock [1 ae Greene]. Peppergrass; 
winter annual.SC:8842;common, rocky slopes. 

sae gordonii (A. Gray) S. Watson var. 

i, Bladder oe winter annual. SC 


Penne micrantha ne Gray) Nieuwl. [Thelypo- 

anthum (A. Gray) S. Watson]. An- 

nual or am SC: 9510, 9645; rare, north- 
facing slopes, Coal Mine Canyon 

Pennellia robinsonii Rollins. Annual or biennial. 

SR: 8256, 8307, 9567, 9597, occasional, grass- 

lands, springs 

ales nasturtium-aquaticum (L.) Hayek.Water- 

DEO, aqu Halle Dla perennial. 

n streams. SR: 9/60; oc- 


Srl springs, Santa Cruz River. 

Schoenocrambe linearifolia (A. Gray) Rollins 
[Sisymbrium linearifolium (A. Gray) Payson]. 
Herbaceous perennial. SC: 8379, 8476; com- 
mon, rocky slopes. SR: 87271; uncommon, 
grasslands 

bes pie (L.) Rollins. Winter annual. SR: 

9; uncommon, sacaton flats. 

a, irio L.London rocket; winter annual. 
SC:9013;common, particularly in overgrazed 
mesquite grasslands. SR: 9046; occasional, 
disturbed areas. 

Streptanthus carinatus C.Wright ex A. Gray ssp. 
arizonicus Weise) Kruckeberg, Rodman 

on]. Twist 


arizonicus S.Wats 


flower; winter annual. SC: 9075, 9090; rare, 
desert grasslands 

Thlaspi montanum L. var. fendleri (A. Gray 
Holmgren [T7.fendleri A.Gray]. oe candytuft; 
herbaceous perennial.SR:8947, uncommon, 


= 
ae) 


north-facing slope adjacent to ae Spring. 
Thysanocarpus curvipes Hook. Lace pod; winter 

annual. SC:8 |, rocky slopes. 
Gactaceae 


\Rritton & Roce in N 


phantha vivipara (Nutt. 

i" Britton & A.Brown [C. vivipara var.bisbeeana 

(Orcutt) L.D. Benson]. Beehive cactus; small 

stem succulent. SC: uncommon, grasslands 
Oc 


casional, grasslands 


Cylindropuntia le is (DC.) FM. Knuth in C. 


Backeberg & F.M.Knuth [O; ial li 
DC]. Desert Christmas cactus, st aecilent SC: 
rare, included based on a photo by R. 


Cyli rerun ee ano Nite M. Kunth i in 
keber 


unt 


ae ree Cane cholla, Holle large 
succulent. SC: common, grasslands and 
rocky slopes. SR: occasional, grasslands, 
Parker Canyon 

Cylindropuntia versicolor (Engelm. ex J. M. 
Coulter) FM. Knuth in C. Backeberg & F. M. 
Kunth [Opuntia versicolor Engelm. ex J.M. 
Coulter]. Staghorn cholla, cholla, siviri; large 
succulent. SC: rare, a single, sterile plant ina 
drainage on the north end of the park ap- 

e this species. 

Echinocereus fendleri (Engelm.) Sencke ex J.N. 
Haage. Hedgehog cactus;small stem succu- 
lent. SC: occasional, grasslands and rocky 
slopes. 

Echinocereus — idissimus (Engelm.) Engelm. ex 
Ha ectinatus (Scheidw.) Engelm. var. 

rigidissimus is (enalan) Engelm. ex Rumpler]. 

ainbow cactus; small stem succulent. SC: 
occasional, rocky slopes. SR: uncommon, 
grasslands. 

Echinomastus intertextus (Engelm.) Britton & 
Rose]. Small stem succulent. SR: 8953; locally 

slands 


pall 


common gras 

Ferocactus wislizeni (Engelm.) Britton & Rose. 
Barrel cactus; large stem succulent. SC: oc- 
casional, grasslands 

Mammillaria grahamii Engelm. [M. microcarpa 
Engelm.]. Fishhook cactus; small stem suc- 
culent. SC: occasional, rocky slopes 

Mammillaria macdougalli Rose in L.H. Bailey]. 
Small, flat succulent. SC: occasional, rocky 
slopes, canyons 

Opuntia engelmannii Salm-Dyck ex Engelm. var 
engelmannii.Engelmann prickly pear, nopal; 
eae parca common, mostly in mes- 

grasslands 


Ba oedl ds. 
ae macrorhiza Engelm. [O. plumbea Rose]. 
ckly pear; small succulent. SR: occasional 
grasslands. 
Opuntia phaeacantha Engelm. Prickly pear, no- 
pal; sprawling succulent. SC: occasional, 
grasslands. SR: uncommon, grasslands. 


684 


Opuntia santa-rita (Griffiths & Hare) Rose. Santa 


Rita prickly pear, nopal; large succulent. SC: 


abundant on rocky slopes 


Campanulaceae 
Lobelia os nalis L. Cardinal flower; perennial 
erb. SC: 8490; uncommon, wet soil along 
streams 
Lobelia fenestralis Cav. Summer annual. SR: 8555; 
incommon, grasslands, sacaton flat 
Nemacladus lees liferus Jepson var. ori Sais 
Thread plant; winter annual. SC: 
9070, 9135; or common, rocky slopes, 


canyons, 

Triodanis holzingeri McVaugh. Venus ane 
Ss; annual. SC: 9359; rare, canyon 

a perfoliata (L.) Nieuwl. rane looking 
ss; annual. SC: 9203, 9223; occasional, can- 
yons. 


oO | 
sav) 


Capp 

Cleome lutea Hook. var. jonesii J.F. Macbride [C. 
jonesil J.F. Macbride]. Bee plant; robust sum- 
mer annual. a soils 
adjacent to Sonoita 

Polanisia Te dra (L.) De ssp. trachysperma 
(Torr.& A. Gray) Iltis [PR trachysperma Torr. & A. 
Gray]. Clammy weed; summer annual. SC: 
8464; occasional, along streams. SR: 8717 
common, Parker Canyon. 


Caprifolia 

| en mexicana K. Presl ex DC. Mexican el- 
der, tdpiro; small tree. SC: 8915; common 
along Sonoita Creek. SR: 9440; occasional, 
Santa Cruz River floodplain. Kartesz (1999) 
includes this taxon as a synonym of the 
widespread S. nigra L. ssp. canadensis (L.) R. 
Bolli. Mexican elder is a low-elevation, early 
spring-flowering species, while the variety 
of S. nigra [S. caerulea Raf.] found in Arizona 
occurs at much higher elevations and flow- 
ers in the summer and fall. 


— 


coy 


Caryophyllaceae 
Cerastium texanum Britton. Mouse-ear chick- 
weed; annual. SC: 9020; rare, ee: 
slopes, Fresno and Coal Mine canyons. 
a molluginea (Lag.) Didr. [D. sperguloides 
Gray]. Drymary; summer annual. SC: 8472; 
mmon, rocky upland 


uNnCcOo 


Silene antirrhina L. Sleepy catchfly; winter annual. 
: 9027; common, rocky slopes. SR: 9307; 


BRIT.ORG/SIDA 22(1) 


uncommon, oak woodland adjacent to 
Parker Canyon. 


Chenopodiaceae 
Atriplex canescens (Pursh) Nutt. var. canescens. 
Saltbush, orache;shrub. SC 
overgrazed ridgetops. 
Atriplex elegans (Moq.) D. Dietr. var.elegans. Sum- 
annual. SR: 9513; occasional, grasslands 
roadsides. 
Atriplex wrightit S. sae Wright saltbush; sum- 
nnual. SR: 8277, 9560; ee Santa 
Cruz ne 
*Chenopodium Giro ONaE: : Goosefoo 
hipasote;annual.SC:8 


ee roadsi 


Sonoita Creek. 

Chenopodium berlandieri Moq. var. sinuatum 
(Murr) Wahl. Pitseed goosefoot; summer an- 
nual. SR: 8704, 8732, 9615; occasional, Parker 
Canyon, grasslan ds 

Chenopodium fremontii S.Watson var. fremontii. 
Go ual. SC:8382; occasional, can 
yons. SR: 9690; uncommon, Parker in. 

lisa neomexicanum Standley var. 
n xicanum. Goosefoot;summer annual. 
SC: a 9548, 9670; Common, Canyons, up- 
lands. SR: 9667: uncommon, below oaks. 

Chenop ? watsoni A. Nelson. nen 
ee ae ar- 
eas, grasslands 

Monolepis nuttalliana (J.A. Schultes) Greene. 
a winter annual. SR: 8941; locally abun 
dant, grasslands. 

*Salsola tragus L.[S. kali L. var. tenuifolia Tausch]. 
Tumbleweed, Russian thistle; summer an- 
nual, SC:uncommon, disturbed sites. SR: 9564; 
abundant, roadsides 


Convolvulaceae 
Convolvulus equitans Benth. [C. incanus Vahl. 
Hoary disc dea ipeae perennial vine. 
C: 9585; uncommon, mostly in canyons. SR: 
8710;0 paces grasslands 
Evolvulus a sinoides (L.) L. var. angustifolia Torr. 
H ous perennial. SC: 9127; rare, rocky 


Hes 
Evolvulus arizonicus ae Gray. MGI BRESOUS peren- 
nial.SC:8575, | canyons, rocky 
slopes. SR: 8308; occasional, grasslands. 
Evolvulus sericeus Sw. var. sericeus. Herbaceous 
ial. SC: 9509; rare, canyons I; 
ae ee gion oak woodenes 


MCLAUGHLIN, FLOR 


AREA AND CAN RAEFCAEI STATE PARK 685 


Ipomoea barbatisepala A. Gray. Morning glory; 
summer annual vine. SC: 8394, 9582; com- 
mon, canyons 

Ipomoea capillacea (Kunth) G. Don [/. muricata 
Cav. sensu Kearney & Peebles (1960)]. Morn- 
ing glory; herbaceous perennial. SR: 9520; 
uncommon, grasslands 


Ipomoea costellata Torr.M 


Cucurbita foetidissima Kunth. Buffalo gourd, 
calabacilla; herbaceous perennial vine. SR: 
456; occasional, grasslands 
Echinopepon wrightii (A. Gray) S. Watson. Wild 
balsam apple; herbaceous annual vine. SC: 
8383; uncommon, canyons 


Cuscutaceae 


annual vine. oS eg 851 . common, can 
sand uy 07, occasional, 
grasslands, Parker Canyon. 

Ipomoea cristulata Hallier f. [ coccinea L. sensu 
Kearney and Peebles (1960)].Star glory, trom- 
pillo;sumn er annual vine. SC:8466;common, 
canyons. SR: 8593; common, Parker Canyon. 

Ipomoea hederacea oo ee ae suM- 

mer annual vine.SC:9 ns 
SR:8595, 9610,9727;common, arn 

Ipomoea longifolia Benth. Morning glory; long- 
stemmed ae SR: 9515;common, 
Oak sav 

ales purpurea .) Roth [/ hirsutula Jacq. f]. 

Tall morning 

8376; occasional, canyons 

? av. var. leptotoma (Torr) J.A. 

~ McDonald [ [/. leptotoma Torr].Morning glory; 
summer perennial vine. SC: 8365; common, 
canyons 

Ipomoea thurberi A. Gray. Morning glory; sum- 

er perennial vine. SC: 8473; rare, canyons. 


fa) 


nnual vine. SC: 


CHMHIOHG 


Crassulaceae 
Crassula connata (Ruiz & Pavén) Berger var. 
ta [Tillaea erecta Hook. & Arn]. Pigmy 
weed; winter annual. SC: 8937; uncommon, 
grasslands 
Sedum cockerellii Britton. Stonecrop; succulent 
perennial herb. SC: 9704; rare, rock walls. 


Cucurbitaceae 
es undulata A.Gray.Meldn loco,melén 
de coyote; DelPadcous Peele) vine. oo 


8561; occasional, grasslands. 
*Citrullus lanatus (Thunb.) Matsum. & Nakai 
Watermelon; annual. SR: 9737; rare, Parker 


Can 
Cucurbi : ee A. Gray. FINget: leaved gourd, 


calabacilla, chichicayota 


nial vine. SC: uncommon, washes aR Unedate 
mon, grasslands. 


Cuscuta erosa Yuncker. Dodder; parasitic vine. SC: 
8504, 8526; uncommon, found on Tragia 
nepetifolia. 

Ericaceae 


i] AA : | | 


SR: 9055; rare,oak savanna.One collection from 
near the north (unfenced) park boundary. 


Euphorbiaceae 

Acalypha neomexicana Muell.-Arg. New Mexican 
ie summer annual. SC: 8389; un- 

on, canyons. SR: 8248, 9590; common, 
grass sland S. 

Acalypha phleoides Cav. [A. lindheimeri Muell.- 
Arg.]. Herbaceous perennial. SR: 8242; occa- 
sional, wash 

Acalypha esmole Riddell. Hornbeam three- 
seeded mercury; summer annual. SC ; 
uncommon, canyons and slopes. SR: 8245; 
occasional, washes. 

Argythamnia neomexicana Muell.-Arg. [Ditaxis 
neomexicana (Muell.-Arg.) Heller]. Herba- 
ceous perennial. SC:8458,8835; uncommon, 
rocky slope 

Chamaesyce abomerennata (Torr. & A. Gray) 
Small [Euphorbia albomarginata Torr.& Gray]. 
ee TS herbaceous eos 

sslands. SR:8243; oc- 


ne disturbed a 
Chamaesyce arizonica ae Arthur (Euphor- 
bia arizonica Engelm.]. Spurge, golondrina, 
herbaceous perennial. SC: 8377, 8412; occa- 
sional, rocky slope 
Chamaesyce capitellata (Engelm.) Millso. [Eu- 
phorbia capitellata Engelm.]. Spurge, 
golondrina; herbaceous Salle SR: 8340, 
8565; occasional, grasslan 
Chamaesyce dioica (Kunth) ie [Euphorbia 
indivisa (Engelm.) Tidestrom]. se 
ae summer annual. SC: 8463; com- 
watercourses. SR: 8332; a 
ee 


pas 


Chamaesyce florida (Engelm.) Millsp. [Euphorbia 

florida Englem.].Golondrina;summer annual. 

common,grasslands and uplands. 

Chamaesyce hirta (L.) Millsp. [Euphorbia hirta LJ. 

Spurge, golondrina; summer annual. SC:8349, 

occasional, canyons. SR: 8587; occasional, 
washes. 


Chamaesyce hyssopifolia (L.) Small oe 
hyssopifolia L.].Golondrina, summer annual. 
SC: 8387; uncommon, canyons. SR: a ,OC- 
casional, washes, grasslands, roadsides 

Chamaesyce melanadenia (Torr) Millsp. [Euphor- 
bia melanadenia Torr]. Spurge, golondrina; 
herbaceous perennial. SC: 8832, 9415; occa- 
sional, rock walls. 

Chamaesyce micramera (Boiss. ex Engelm.) 
Wooton & Standley [Euphorbia micromera 
Boiss.]. pulse golondrina; summer annual. 
SC: 8661; uncommon, drainages. 

Chamaesyce pediculifera (Engelm.) Rose & 
Standley var. pediculifera [Euphorbia 
pediculifera Engelm.]. Spurge, golondrina; 
herbaceous perennial. SC: 8359; occasional, 
rocky slopes. 

Chamaesyce revoluta (Engelm.) Small [Euphor- 
bia revoluta Engelm.]. Golondrina; summer 
annual. SC: 8521; rare, rocky slopes. SR: 8716; 
rare, Parker Canyon 

Chamaesyce serpyllifolia (Pers.) Small [Euphorbia 
sea i Sy coal ee summer 

nnual.SR:8 8695 i | sslands 

Chamaesyce ne (Engelm. ex di cite eX 
Parish [Euphorbia setiloba Engelm.]. Spurge, 
golondrina; summer annual. SC: 86/0; rare, 
along Sonoita Creek. 

Chamaesyce stictospora (Engelm.) Small [Eu- 
horbia stictospora Engelm.]. Spurge, 
golondrina, summer annnal. SR: 8321, 8600; 
occasional, grasslands. 

Chamaesyce vermiculata (Raf.) House [Euphorbia 
vermiculata Raf]. Spurge, golondrina; sum- 

mer annual. SR: 8304; occasional, springs 

Cnidoscolus angustidens Torr. Mala mujer, 
sla a; Liles perennial. SC: 8456; 


common, S. 
Croton ae halen ce ee [C. 
L / | ] | | Jef | 


ceous perennial. SC: 9542 ional, rocky 
slopes. SR: 8601 - occasional, grasslands. 
Croton texensis (Klotzsch) Muell.-Arg. var. texensis. 


BRIT.ORG/SIDA 22(1) 


Dove weed, yerba de la tértola, summer an- 
nual. SC: 8668, 8692; uncommon, sce soils 
along Sonoita Creek. S i, 
Santa Cruz River patton. 

Euphorbia bilobata Engelm.Summer annual. SR: 
9613, 9718; uncommon, Parker Canyon. 

Euphorbia chamaesula Boiss. Herbaceous peren- 
nial. SR: 8281, 8333, 9312; occasional, grass- 
lands, roadsides. 

ss cuphosperma (Engelm.) Boiss. [E. 
dentata Michx. var. cuphosperma (Engelm.) 
Fein Painted ae summer annual. SC: 

nds 


ee cae fae Spurge; summer 
annual. SC: ee uncommon, mesquite 
shes 


Euphorbia ae ee bainted ae sum- 
mer annual. SC: 8350; uncommon, canyons 
and uplands.SR:8377;occasional, grasslands 


springs. 

Euphorbia radians Benth. Painted spurge; herba- 
ceous perennial. SR: 9/09; common, grass- 
lands. 

Jatropha macrorhiza Benth. var. septemfida 
Engelm. Herbaceous perennial. SC: 9472; 
uncommon, canyons and uplan S 

Manihot angustiloba (Torr) Muell.-Arg. Herba- 

ous perennial. SC: 9547; rare, canyons. 
or ens polygonoides Nutt. ex Spreng. Her- 
baceous perennial. SC: 8488; rare, Fresno 
anyon. 


Tragia laciniata (Torr.) Muell.-Arg. Nose burn; 
h ous perennial. SR: 8379; occasional, 
a on north-facing slopes. 
Tragia het Cav. var. dissecta Muell.-Arg. 
se burn; herbaceous perennial. SC: 8358, 
8500; eae rocky slopes. 


Fabaceae 
Acacia angustissima (P. Mill.) Kuntze. White ball 
acacia; oe a ial 8; occasional, ridge 
crests. SR: | 
Plants appearing to belong to different va- 
rieties may occur mixed in the same popu- 
lations. 
Acacia greggii A. Gray. Catclaw, una o_o 
ae os coe Fonlbae ca 
n.Large ae ee 8503, 


grasslands 


9492; rare, aoe 
Amorpha fruticosa L. ao ae large shrub. SC: 
9286; rare, one patch found in Fresno Canyon. 


MCLAUGHLIN, FLORA OF 


AREA AND SAN RAFAEL STATE PARK 687 


Astragalus allochrous A. Gray var. allochrous. 
Halfmoon loco; winter annual. SC: 9730; un- 
common, along Sonoita Creek. 

Astragalus arizonicus A. Gray. Locoweed; herba- 
Cc s perennial. SC: 9729, 9219; common, 
uplands. 

ee humistratus A. Gray var. sonorae (A. 

rdy 


Astragalus nothoxys A. Gray. Locoweed; herba- 
ceous perennial. SC: 9237; uncommon, can- 
yons and uplands. SR: 9306; common, grass- 
lands. 

en nuttallianus DC. var. tri a. Torr. 

ocoweed; winter annual. S 04; 
sundae tae SR:9093; sbuneants grass- 
lan 


ae pinia gilliesii (Hook.) Wallich ex D. Dietr. 
Bird-of- Ghee ale 9473; rare, gallery 
he along Sonoita Cre 
Call eee eriophylla ey var. eriophylla. Fairy 
duster;low shrub.$C:932/,occasional, uplands. 
Calli aa humilis Benth. var. reticulata (A. Gray) 
n [C. reticulata A. Gray]. False mes- 
ones perennial a 8559: occa- 
ands 


Chamaecrista nictitans (L.) Moench ssp. nictitans 
var. leptadenia (Greenman) Gandhi & Hatch 
[Cassia leptadenia Greenman]. Yerba del piojo; 
summer annual. SC: 8403; occasional, grass- 
lands and uplands. SR: 8257; occasional, 
grasslands. 

Cologania angustifolia Kunth [C. longifolia A. 
Gray]. Herbaceous perennial. SC: 9682; rare, 
deel ae SR: E.A. Lewis 004; uncom- 

ker Cany 

ee ane ee Lavin var. caribaea 

a edwardsii A. Gray]. Herbaceous pe- 
Ae rae ie ne on north- 
facing slopes. 

Crotalaria js Ortega. Rattle box; summer 

|. SC: 7, occasional, canyons and 
Ge oie in Parker 
Canyon. 


Dalea albiflora A.Gray. Herbaceous perennial.SC: 
uncommon Sey aes SR: 8507, 8579; 


baa ae eae ex ie var. oligophylla 
rr.) Shinners [Petalostemon candidum 
ae Michx. var. oligophyllum (Torr.) Her- 


= 


mann].White prairie clover; herbaceous pe- 
rennial. SR: 9390; occasional, grasslands. 

Dalea formosa Torr. Feather plume, shrub. SC: 
8674,9 9068; locally common uplands 

Dalea grayi (Vail) L.O. Williams. Herbaceous pe- 
rennial. SR: 8506; uncommon, grasslands. 

Dalea lachnostachys A.Gray. Herbaceous peren- 
nial. SC: 9322; uncommon, oi uplands. SR: 
8722; uncommon, gras cs 

Dalea leporina (Ait.) niece ee dalea; sum- 
mer annual. SR:8620;uncommon, grasslands. 

Dalea nana Torr. ex A. Gray var. carnescens Kear- 
ney & Peebles. Herbaceous perennial. SC: 
8524, 9287, occasional, rocky uplands. SR: 
8549C; occasional, grasslands 

Dalea neomexicana (A. Gray) A. Gray var. 
neomexicana. Herbaceous perennial. SC: 
8682, 9273; uncommon, rocky uplands. 

Dalea pogonathera A. Gray var. pogonathera. 
Yerba del coraz6n; herbaceous perennial. SC: 
8512?:0c 


casional, uplands 

Dalea pringlei A.Gray. Herbaceous perennial. SC: 
8752, 9222, 9288; occasional, canyon, rocky 

slo 

Dalea ee H.S. Gentry. Indigo bush, shrub. 
SC: 8839, 9126; occasional, canyons, rocky 
slopes. 

Dalea versicolor Zucc. ssp. versicolor var. sessilis 

A. Gray) BanSDy ingige bush; shrub. SR: 

9305, rare, oal d adjacent to Parker 
Canyon. 

Desmanthus cooleyi (Eaton) Trel. Herbaceous 
perennial.SC: 957 T;rare, mesquite grasslands. 
SR: 9529, E.A. Lewis 002; common, grasslands 
and roadsides. 
maditim bor 


qulon A. Gray. Tick clover, her- 
baceous perennial. SC: 8489; rare, canyon 

Desmodium neomexicanum A. Gray. Tick an 
summer annual. SC: 8541; rare, en SR: 

Parker C 

Desnouilinn ee (P. ai, Hitche. var. 
exiguum (A. Gray) Schub. Tick clover, summer 
annual. SC: 8681, 8687; rare, canyons 

y. Tick clover, ro- 

bust hetbacsous eer SC: 8436, 8443, 

9553, 9701; common, canyons and rocky 


Ooditym nsilocarnism A. Gra 
tf 


slopes. 
ee rosei Schub. Tick clover; summer 
annual. SR: 9677; uncommon, Parker Canyon. 
Erythrina flabelliformis Kearney. Southwestern 


688 


coralbean, chilicote, pionilla; shrub. SC: 9411; 


common, rocky slopes. 

Eysenhardtia orthocarpa (A. Gray) S. Watson [E. 
polystachya (Ortega) Sarg. sensu Kearney & 
Peebles (1960)]. Kidneywood, palo dulce; 
large shrub. SC: 84317, 9412; common, rocky 


slopes. 

Galactia wrightii A. Gray var. mollissima Kearney 
& Peebles. Herbaceous perennial vine. SC: 
8373; occasional, canyons. SR: 9459; rare, 
grasslands. 

Indigofera sphaerocarpa A.Gray.|ndigo, shrub. SC: 

orth-facing slope, Coal Mine Canyon. 

*Lotus aoe sel Bicasieot trefoil; herba- 

sprin 


Note River botto 
me greenei Ottley ex ae & Peebles. Deer 
vetch. 9022. Herbaceous perennial; uncom- 
mon, oak savannas. SR:8949 


grasslands. 

Lotus humistratus Greene. Winter annual. SC: 
8 ommon, es coals grasslands. SR: 

| | grasslands 

Lupinus brevicaulis S. ia Short-stemmed 
lupine; winter annual. SC: 9080; occasional, 
mostly along drainages. SR: 9/05; common, 
grasslands. 

Lupinus concinnus J.G. Agardh. ssp. orcuttil . 
Watson) D. Dunn. Elegant lupine; winte 
nual.SC:8930, 9063; occasional, mostly fone 
drainages. SR: 9107; occasional, Parker 


Canyon. 

Lupinus sparsiflora Benth. ssp. mohavensis 
Dzie cen & D. Dunn. Lupine; winter an- 

08; locally abundant, slopes. 

Macroptil paae ce (Ortega) Delgado 
[Phaseolus heterophyllus Willd.]. Herbaceous 
perennial vine. SC: 8479; uncommon, can 
yons. SR: 9517, 9569, 9659; common, grass- 
lands 

Marina calycosa (A. Gray) Barneby [Dalea 
calycosa A. Gray]. Annual. SC: 8679, 9274; 
uncommon, uplands. SR: 9382; uncommon, 
oak savannas 

*Medicago lupulina L. Black medic; annual. SR: 
9313, 9386; common, springs, Santa Cruz 
River bottom. 

lea sativa L. Alfalfa; se perennial. 
SR: side 


*Meli fe indicus (L.) All. one ion sweet 


BRIT.ORG/SIDA 22(1) 


clover, alfalfilla; annual. SC: 9265; occasional 
along Sonoita Creek. SR: 9298; occasional, 
Santa Cruz River floodplain. 

*Melilotus officinalis (L.) Lam. Sweet clover; tall 
annual. SR 8790, 9426; both white and yel- 
low-flowered forms abundant, Santa Cruz 
River bottom 

Mimosa aculeaticarpa Ortega var. biuncifera 

Benth.) Barneby [M. biuncifera Benth.].Wait- 

a-minute, cat's claw; shru > : 9470; com- 
mon, rocky slopes, canyons. SR:| llyab 
dant, lastee 

Ai Benth. Shrub. SC: 9353: com- 

mon, rocky slopes. SR:8337,occasional, grass- 
ands. 


— 


| 
Nissolia schottii (Torr.) A. Gray. Herbaceous pe- 
nnial vine. SC:8409; uncommon, north- fac- 


ing slop 
Phaseolus Stee A. Gray var. acut uae 
ep >summer annual. SC: 9639; un- 
ommon, oe SR: 9616; uncommon, 
Parker Canyon 
Prosopis velutina W [Pjuliflora(S DC. 
var. velutina (Wooton) Sarg.].Velvet mesquite 
mezquite; small tree. SC: 9587; common, 
washes, ridgetops, gentle slopes. SR: uncom- 
mon, grasslands. 
eAIGHGIE ium a aOR a (Pursh) Rydberg 
[Psoralea tenuiflora Pursh]. Scurf pea; herba- 
ceous perennial. SC: 9287; rare, canyons. SR: 
939 | {land adj to 


Parker Canyon. 

Rhynchosia minima (L.) DC. Rosary bean; herba- 
eous perennial vine. SC: 9706, 9733; rare, 
rocky slopes, canyons 

Rhynchosia senna Gillies ex Hook. var. texana 
(Torr.& A.Gray) M.C. Johnston [R. texana Torr. 
& A. Gray]. Rosary bean; herbaceous peren- 
nial vine. SC: uncommon, north-facing 
slopes, canyons. SR: 8558, 9580; occasional, 
grasslands. 

Senna bauhinioides (A. Gray) Irwin & Barneby 
[Cassia bauhinioides A. Gray]. Herbaceous 
perennial. SC: 9325; uncommon, uplands 

Senna hirsuta (L.) Irwin & Barneby var.glaberrima 
M.E. Jones) Irwin & Barneby [Cassia 
leptocarpa flees Robust herbaceous pe- 
rennial. SC: 8381, 8465; uncommon, canyon 
bottom. SR: uncommon, grasslands. 

Sphinctospermum constrictum (S.Watson) Rose. 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL 


AREA AND SAN RAFAFL STATE PARK 689 


Summer annual. SC: 8460; rare, mesquite 
grasslands. 

Tephrosia tenella A. Gray. Summer annual. SC: 
8542; uncommon, a dal SR: 8584; 


unco Parker Can 
Tephrosia ae Schlecht. Herbaeou peren- 
nial.SC:8483; rare, found trop, 


where it was locally abundant. 

*Trifolium fragiferum L. Clover; herbaceous pe- 
rennial. SR: 9437; uncommon, Santa Cruz 
River bottom 

Trifolium mucronatum Willd. ex Spreng. ssp 
lacerum (Greene) J. Gillett [7. lacerum 
Greene]. Clover; herbaceous perennial. SR: 

9523, rare, wet groun 
*Trifolium pratense L. Red clover, eae 


perennial. SR: 9489; a Cruz 


Vicia ludoviciana Nutt. ssp. /[udoviciana. Vetch, 
winter annual vine. SC: 8874; common, 


ashes. 
ee gemella Vogel [Z. diphylla (L.) Pers. sensu 
Kearney & Peebles (1960)]. Herbaceous pe- 


Cp. nez. 


rennial. SR: ip) 7; unc ommon, arasslands 


I ,encino 


Quercus arizonica Sarg. Arizona 
blanco; tree. SR: 9452, 9457, 9663, 9738 com- 
mon in woodland adjacent to Parker Canyon. 

Quercus emoryi Torr.Emory oak, bellota; tree. SC: 
8533; uncommon, canyons, north-facing 
slopes. SR: 9516; common, oak savannas and 
oak woodlands 

ales grisea Liebm. Gray oak; tree. SR: 8344, 

afew trees just east of the 
Santa Cruz River appear to be this species. 
oe ob! ee, Torr. Mexican blue oak, 

8775, occasional, north- 

Hee slopes, canyons. 


oazul tre 


Fouquieriaceae 
Fouquieria splendens Engelm. Coach whip, oco- 
tillo; shrub. SC: uncommon, rocky slopes 


Fumariaceae 

Corydalis curvisiliqua Engelm. ssp. occidentalis 
Engelm. ex A. Gray) W.A. Weber [C. aurea 
Willd. ssp. occidentalis (Englem. ex A. Gray) 
G.B. Ownbey]. Golden corydalis; winter an- 
nual. SC: 8869; common, rocky slopes, can- 
yons. SR: 895 1; occasional, Parker Canyon. 


Garryaceae 
Garrya wrightii Torr. Silk tassel; al shrub, SC: 
;uncommon, ca 


Gentianaceae 
Centaurium calycosum (Buckley) Fernald. 
Buckley's centaury, annual. SC: 9272; uncom- 
alon ; 
Centaurium nudicaule (Engelm.) B. L. Robinson. 
Annual. SC: 9230, 9295; uncommon along 
streams. 


Gereningese 
Erodium cicutarium (L.) L’Her. ex Ait. ssp. 
cicutarium. Filaree, alfilaria, afilerillo; winter 
annual. SC: 9134 ional, mostly in mes- 
quite grasslands. SR: 9050; uncommon, dis- 


rbed areas. 
Erodium texanum A. Gray. Heron bill; winter an- 
nual. SC: 9072; uncommon, canyons and 
mesquite grasslands. 
Grossulariaceae 
Ribes aqureum Pursh var.aureum.Golden current, 
shrub. SR: 9759; rare, Sharp Spring, Santa Cruz 
River bottom. 


Haloragaceae 

Myriophyllum cf. sibiricum Komarov [M. 
exalbescens Fern.].Water milfoil; subrnerged 
aquatic perennial. SR: 9247; rare, Santa Cruz 
River bottom; not found in flower. 


Hydrangeaceae 
Fendlera wrightii (A. Gray) Heller [Ff rupicola A. 
Gray var. wrightii A. Gray]. Large shrub. SC 
8407, 8836, uncommon, north-facing slobes 
anyons. 
Been: microphyllus A.Gray. Mock orange, 
shrub. SC: 9218, 9357; rare, north-facing slope 
in Fresno Canyon. An atypically low-eleva- 
tion record for this species. 


Hydrophyllaceae 
ees buts oe Heller. Winter annual. 
85 canyons, rocky 


fae 
Nama upigdrn A. Gray. eae ae fare ae 
C: 8913 s adjacen 
stre ee 
Phacelia al 5 A. Gray. purole < phacelia;win- 
- 9062, 9207, 9268; Common, 
mostly ae mesquites. 


[ea prewar . 
4: it 


Phacelia arizonica A. Gray.Winter annual. SC: 8856; 
common, ec nes quite eine! pune 
aiercd areas.$ sslands 

Phacelia bycina Wooton & atta Winter 
annual. SC: 8905, 9206; occasional, rocky 
slopes. 

Phacelia caerulea Greene.Winter annual. SC:897 7 

nal, grasslands, c 

ha ee eve ae winter 
annual. SC: 9071; locally abundant, rocky 

slopes. 

Phacelia sonoitensis S.P. McLaughlin ined. Win- 
ter annual. SC: 9088, 9123,9206, 9226; uncom- 
ene slopes. 


Hypericaceae 

Hypericum scouleri Hook. ssp. scouleri [H. 
formosum Kunth]. Herbaceous perennial. SR: 
9577, rare, SDrings. 

Juglandaceae 

Juglans major (Torr.) lee Arizona walnut, 
nogal; large tree. SC: 9 |,along 
eG Creek and in the larger canyons. SR: 

‘ , Parker Canyon. 


Krameriaceae 

Krameria erecta Willd.ex JA on ites [K. parvifolia 
Benth.]. Ratany, tamichi; low hemiparasitic 
shrub, SC: 9227, pe rocky slopes. SR: 
9380; uncommon, oak savannas. 

Krameria lanceolata Torr. Ratany; hemiparasitic 

herbaceou | ennial SR:9393 


grasslands, 


Lamiaceae 
Hedeoma dentata Torr.Mock pennyroyal;herba- 
ceous Hees SC: 851]; uncommon, can- 
yons. SR: 8727; uncommon, oak woodland 
adjacent to Parker Can 
Hedeoma oblongifolia (A.Gray) Heller Mock pen- 
yroyal; herbaceous perennial. SC: 9584; 
uncommon, canyons 
*Lamium amplexicaule L. Dead nettle; winter 
annual. SC: 8970; uncommon, mostly along 


on 


Sonoita Creek. 

*Marrubium vulgare L. Common horehound, 
marrubio; herbaceous persis) alt 8326, 
occasional, grasslands 
sist around abandoned homesteads. 

Monarda citriodora Cerv.ex Lag. ssp. austromon- 

a (Epling) Scora [M. austromontana 
Epling]. Bee balm, horsemint, orégano; her- 


BRIT.ORG/SIDA 22(1) 


baceous perennial. SR: 8238; occasional, 
grasslands. 

Prunella vulgaris L. ssp. lanceolatus (W. Bart.) 
Hultén. Self-heal; herbaceous perennial. SR: 
9522, rare, wet ground. 

Salvi peal i A. Gray. Sage; low shrub. SC: 8683, 
,Canyons 
Salvia ree Benth.Summer annual. SC: 8398, 
1; occasional, canyons and uplands. SR: 
82 a occasional, grassland. 

Stachys coccinea Ortega. Texas betony; herba- 
ceous perennial. SC: 8872; locally common 
in canyons 

Trichostema arizonicum A. Gray. Blue curls; her- 
baceous perennial. SC: 8680, 9546; rare, 
canyons 


Linaceae 
Linum deseo (Engelm.) Heller. Plains flax; 
a al.SC: 9140, 9278; uncommon, uplands 


-§317-uncommon. arasslands 


Loasaceae 

Menitzelia albicaulis (Douglas ex Hook.) Douglas 
ex Torr.& A.Gray. Small flowered blazing star; 
winter annual. SC: 8847, 885 7;occasional, rocky 
slopes. SR: 9098; occasional, Parker Canyon. 

alee isolata H.S.Gentry [M. asperula Wooton 

tandley sensu Kearney and Peebles 

ae ure ee 
8404; occasional, rocky slopes 

Mentzelia multiflora (Nutt.) A. Gray. Adonis blaz- 
ing star, yerba pegajosa; biennial or herba- 
ceous perennial. SC: 8690; occasional, sandy 
soils adjacent to Sonoita Creek. SR: 9373: ac- 
casional, grasslands, roadsides 


Lythraceae 
Lythrum californicum Torr. & A. Gray. Loosestrife, 
yerba del cancer, herbaceous perennial. SR: 
springs 
Rotala ramosior (L.) Koehne. Toothe cup; sum- 
mer annual. SC: 9674; rare, wet soil of canyon 
bottom 


Malpighiac 
PaPICa DS en a Rich. Herbaceous perennial. SR: 
oak savannas, grasslands 


Janusia gracilis A. Gray. Twining shrub. SC: 8516 
rare, ridgetops 

Malvaceae 

Abutilon incanum (Link) Sweet. Indian mallow, 


MCLAUGHLIN, FLORA OF URAL 


AREA AND CAN RALFAFI STATE PARK 691 


pelotazo; small shrub. SC: 8844; un 
rocky slopes. 

Abutilon mollicomum (Willd.) Sweet [A. sonorae 
A. Gray]. Tall herbaceous perennial. SC: 8447; 


= 
= 


unco /canyons. 

Abutilon parvulum A.Gray.Small-leaved abutilon; 
herbaceous perennial. SC: 9556; rare, can- 
yons. 

Anoda abutiloides A. Gray. Tall herbaceous pe- 
rennial.SC:8386, 8442, 8753;occasional, rocky 
slopes and canyons. 

Anoda cristata (L.) Schlecht. Spurred anoda, 

uesito; summer annual. SC: 8462, 8662; oc- 
casional, mostly in canyons. SR: 8566; occa- 
sional, grasslands. 

Gossypium thurberi Todaro. Desert cotton, shrub. 
SC: 8414; locally common, canyons, slopes, 
ridgetops. 

Herissantia crispa (L.) Brizicky [Gayoides crispum 
(L.) Small]. Herbaceous perennial. SC: 8477 


8517; uncommon, rocky slopes and 
ridgetops 

Hit [ S.Watson.R llow;herb 
ceous perennial. SC:8424 rocky 


slopes. 
*Malva ees L.Cheese weed; winter annual. 
SR: common, disturbed areas 
eee tan (A. Gray) Fryxell [Sida 
physocalyx A. Gray]. Herbaceous ie 
tie ncommon, canyons. SR: 8589; un- 
ker Canyon. 
sa cut P. Mill. [S. procumbens Sw.]. Pros- 
herbaceous perennial. SC: 8546; occa- 
Bee ede 8 oe eee 
lands. 
ae neomexicana A.Gray. Herbaceous perennial. 
580; occasional, Tee | 
a pigs L. Prickly mallow; summer annual. 
R: 8336, 8568; aes grasslan 
eres angustifolia (Cav.) G. Don var. 
cuspidata (A. Gray) Kearney. Narrow-leaved 
globe mallow; herbaceous perennial. SR: 
8552; occasional, mostly below oaks. 
Sphaeralcea fendleri A. Gray var. fendleri. Globe 
mallow; robust herbaceous perennial. SR: 
9687; uncommon, Santa Cruz River flood- 
plain. 
Sphaeralcea laxa Wooton & Standley. Caliche 
globe mallow; herbaceous perennial. SC: 
8811; occasional, rocky slopes 


Menispermaceae 
Cl Hf ait ital) 


is DC. Snail seed, correhuela; 
woody vine. SC: 9478; uncommon, canyons, 
rocky slopes. 


uginaceae 
*Mollugo verticillata L.lndian chickweed; annual. 
SC: 8413; uncommon, rocky slopes. SR: 8288; 
common grasslands 


Moraceae 

*Morus alba L. Mulberry, mora. Tree. SF: rare, a 
single plant persisting near ranch buildings. 

Morus ee EUKICY. Texas ee mora; 
small tre / anyons and 
rocky se 


Nyctaginaceae 
Allionia incarnata L. Trailing four o' clock, wind- 
ne sone herbaceous ee SC:8455, 


] (0) 
ee coccinea i Mi Red ering ag 


srssonds. 5 SR: a 1; occasional, gras ae 
disturbed a 
Boerhavia erecta fe Summer annual. SC: 8397, 
10; uncommon, canyons and uplands. SR: 
592; occasional, Parker Canyon 
Boerhavia intermedia M.E. Jones. ace -winged 
ringstern; summer annual. SC: 8514; occa- 
sional, mesquite grasslan 
Boerhavia megaptera Standley. ae annual. 
5- rare, grasslands 
Boerhavia purpurascens A.Gray. Purple spiderling; 
summer annual. SC: 8522; occasional, rocky 
slopes. SR:8775; uncommon, n, Pa rker Canyon. 
Boerhavia scandens L. [Commicarpus scandens 
(L.) Standley]. Herbaceous eens 
mon,below trees and shrubs in canyons. 
eee aG spicata ule Summer annual. SC: 
|, sandy soil of canyon bot- 
tom. SR: 8714, 9715; uncommon, Parker Can- 


Mirabilis coccinea (Torr.) Benth. & Hook. f. 
[Oxybaphus coccineus Torr].Four o'clock, her- 
baceous ae SC: 9354: rare, Fresno Can- 
yon. SR: 9379, sslands 

Mirabilis eee (Pursh) Heimer| ae 
linearis (Pursh) B. L. Robinson]. Herbaceous 
perennial. SR: 8587, 8709; uncommon, grass- 


lands. 
Mirabilis longiflora L. var. wrightiana (A. Gray ex 


Britton & Kearney) Kearney & Peebles. Sweet 
four o'clock, maravilla; herbaceous perennial. 

SC: 9505; rare, north-facing slopes. SR: 8730, 
9620; uncommon, oak savannas. 


Oleaceae 

Fraxinus velutina Torr. Velvet ash, fresno; tree. SC: 
8399; common along Sonoita Creek and in 
larger canyons. SR: 8735; uncommon, Parker 
Canyon 

baie sabia A. Gray. Small shrub. SC: 8675, 


8,932 4; locally common cany 


Onagraceae 
Camissonia californica (Nutt. ex Torr. & A. Gray 
en [Oenothera leptocarpa Greene}. Win- 

ter en SC: 9009, 9122; occasional, can- 
yons and rocky slopes 

Camissonia chamaenerioides (A. Gray) Raven 
[Oenothera chamaenerioides A.Gray].Winter 
annual. SC: 9 ncommon, canyons 

Camissonia claviformis (Torr. & Frémont) Raven 
ssp. peeblesii (Munz) Raven [Oenothera 
clavaeformis Torr. & Frémont var. peeblesii 
Munz]. Winter annual. SC: 9077; rare, one 
population seen in — of Sonoita Creek. 

Epilobium canum (Greene) Raven ssp. latifolium 
(Hook.) Raven ee ia (Hook.) 

Greene var. arizonica (Davidson) Hilend]. 

Hummingbird trumpet; herbaceous peren- 

nial. SC: 8750; i Ica 


Cnilnh; MNiar Rifle Pi |, W | | 


= 


ae 


Us 3K 
Willow weed; herbaceous perennial. SR:8274; 


common prings 

Gaura hexandra Ortega ssp. gracilis (Wooton & 
Standley) Raven & Gregory [G. gracilis 
Wenn & seat oo one, 
SC:8 ons. SR: 8260, 9481: 


common, cow, areas 
Gaura mollis James [G. parviflora Douglas ex 
Lehm.]. Lizard tail, velvet leaf gaura; tall her- 
baceous perennial. SC: 8654; occasional, 
mostly along Sonoita Creek. SR: 8263; com- 
mon, Sharp Spring, low areas, roadsides. 
Oenothera albicaulis Pursh. Evening primrose; 
annual. SR: 9239, 9477, 9514: locally 
sslands 
oer est Nutt. ssp. marginata (Nutt. 
x Hook. & Arn.) Munz. Evening primrose; 
reacts eine SC:9117; Nae a 
SR: assla 


BRIT.ORG/SIDA 22(1 


= 


Oenothera primiveris A. Gray ssp. primiveris. Yel- 
low desert primrose; herbaceous perennial. 
SC: 8834,8918 asslands. SR:8940; 
common, south-facing slopes. 

Oenothera pubescens Willd. ex Spreng [O. 
laciniata Hill ssp. pubescens]. Evening prim- 
rose;annual. SR: 9378, 9480, 9578:uncommon, 
grasslands. 

aiid rosea L'Her. ex Ait. Sun drops; herba- 

eous perennial. SC: 8655; uncommon, wet 
a along edge of eae Creek. SR: 8278; 


common, springs 


Orobanchaceae 

Orobanche cooper! (A. Gray) Heller ssp. cooperi 
[O. ludoviciana Nut var, cooper! ( (A. Gra 
B herbaceous 


perennial. SC: 9269, 9328; occasional, grass- 
lands, canyons. 
Oxalidaceae 
Oxalis albicans Kunth ssp. albicans. Wood sorrel, 
ee herbaceous perennial. SC: 8746; 
on,mostly along Sonoita Creek. SR: 
ee occasional, grasslands. 
= ions A. Gray [O. amplifolia (Trel.) 
nsu Kearney and Peebles (1960)]. 
ie ne herbaceous perennial.SC: 9540, 
uncommon, delet) ery Ete along 
Sonoita Creek, canyons.S 
Parker Canyon 


?4-UNn HTMTOT, 


Papaveraceae 

Argemone mexicana L. Prickly poppy; annual. SC: 
9290; rare, along old railroad right-of-way 

ong Sonoita Creek, possibly introduced. 

pent pleiacantha Greene . saree 

Prickly poppy; annual. SC: 9 common 

along nae Creek and in canyons SR:9425; 

co oadsides, grasslan 


Eschscholzia californica Cham. ssp. mexicana 

crate Clark [E. Greene]. Mexi- 

an poppy, amapola del campo; winter an- 
ee nds 


Passifloraceae 
Passiflora mexicana Juss. Passion flower, Herba- 
perennial vine. SC:un 
forest, canyons, not found in flower. 


common gallery 


Pedaliaceae 
Proboscidea parviflora (Wooton) Wooton & 
Standley ssp. parviflora. Devil's claw, gatitio; 


MCLAUGHLIN, FLORA OF 


AREA AND CAN RAFAFI STATE PARK 693 


summer annual. SC: 8374 
soils adjacent to cone Creek. a 8335; 
common, grasslands 


Phytolaccaceae 

Rivina humilis L. Pigeon berry, rouge plant, 
coralito; herbaceous perennial. SC: 8362 
commoan, canyons 


Plantaginaceae 
*Plantago major L.common plantain;annual.SC: 
8742, 9314; rare, along Sonoita Cree 
Plantago patagonica Jacq. [P. purshii Roemer & 
Schultes]. Plantain; winter annual. SC: 8927; 
abundant, uplands. SR: 9092;abundant, grass- 
ds 


Plantago virginica L. Pale-seeded plantain; win- 
ter annual. SC: 9225, 9231; uncommon, mostly 
along pce in wet soil. SR: 9244; locally 


PVA RA 


comm 
ine oe and probably native in 
Arizona 


Platanaceae 

Platanus wrightii S. Watson. Arizona sycamore, 
aliso; tree. SC: rare, present as a few saplings 
along Sonoita Creek. 


Plumbaginaceae 

Plumbago scandens L. Leadwort, hierba de 
alacrdn, pitillo; herbaceous perennial. SC: 
8408, uncommon, canyons, rocky 


slopes. 


Polemoniaceae 

pues diffusum (A. Gray) Mason. Winter an- 
nual. SC: 9725, 9143; uncommon, mesquite 
grasslands. SR: 9240; occasional grasslands 

Gilia flavocincta A. Nelson ssp. australis (A. & V. 
Grant) A. Day & V. Grant. Winter annual. SC: 
8860, 9061;common, gallery forests, canyons, 
and uplands. SR: 8942 grasslands 

Gilia mexicana A. & V. Gane Winter annual. SE 
9085;uncommon, mostly ridgetops. SR: 9108; 


common, grasslands. 

Ipomopsis macombii (Torr. ex A. Gray) V. Grant 
[Gilia macoumbii Torr]. Herbaceous peren- 
Ue eee ac ea 


c 


yons. lesen rasslands 


Nel 


Linanthus e ssp.aureus.Win- 
ae SE: 9089:1 rare, a Saab 
sa bigelovii (A. Gray) Greene. Winter 
SC: 8861; uncommon, canyons. 


> 


Phlox gracilis (Hook.) Greene [Microsteris gracilis 
(Hook.) Greene].Winter annual. SC: 9.214; rare 
Fresno Canyon 


Polygalac 
Polygal Hae te Milkwort; herbaceous peren- 
nial. SR: 8560, 8737; uncommon, grasslands. 
ies barbeyana Chod. [P. longa Blake]. Milk- 
t; herbaceous perennial. SC: 9405; un- 
common, canyons, rocky slopes. SR: 95217; 
nd 


uncommon, grassla 


Polygala obscura Benth. Milkwort; herbaceous 
perennial. SC: 8749, 9323; uncommon, can- 
yons, rocky slopes. SR: 9572; uncommon, 
grasslands. 

Polygona 

Eriogonum ae Torr.var.abertianum.Wild 
buckwheat; annual. SC:8384;common,all ter- 
restrial habitats. SR: 8330, 8331; occasional, 

rasslands. 

suena polycladon Benth. Sorrel buckwheat; 

mer annual. SC: 8517 oe sas 
po to eile Creek, other dis- 
turbed are 7,occasional, nen 
aerard ee i Torr. ex Benth. var. wrightii. 
ht buckwheat; low shrub. SC: 9729; oc- 
ae ae rocky slopes. SR: 8779; 
common, grasslands. 

*Polygonum aviculare L. Prostrate knotweed, 
chilillo; herbaceous perennial.SR:8342; locally 
abundant, tanks 

pay lapel ffolium L.Willow smarty 

|.SR:8266; pally 


abundant, cattle Anke Santa Cruz River bot- 


tom. 

Polygonum pensylvanica L. Pinkweed, chilillo; 
herbaceous perennial. cea ie aa 
dant, low areas, cattle ta 

Polygonum punctatum ale Var. - confertiflorum 
(Meisn.) Fassett. Water smartweed, chilillo; 
herbaceous perennial. SC: 8497, 8653; occa- 
sional, stream edges. 

*Rumex crispus L. Curley dock; herbaceous pe- 
rennial. SR: 8780, 9427; occasional, springs. 

Rumex hymenosepalus Torr. Wild rhubarb, 
cahaigre; herbaceous perennial. SC: uncom- 
mon, gallery forest along Sonoita Creek. 

Rumex violescens Rech. f. Herbaceous perennial. 
SC: 9318; uncommon, edge of Sonoita Creek. 


Portulacaceae 

Calandrinia ciliata (Ruiz & Pavén) DC. Red maids; 
winter annual. SC: 8854; common, canyons, 
grasslands. SR: 8938; common, low areas, 
roadsides 

Cistanthe ae (A. Gray) Hershkovitz var. 
arizonica (J. T. Howell) Kartesz & Gandhi 
[Cal ean parryi A. Gray]. Winter annual. 
SC: 8928; uncommon, grasslands 

Portulaca suffrutescens Engelm.Purslane; herba- 
ceous perennial. SC: 8420; common, mes- 
quite grasslands, ridgetops. SR:8567; uncom- 
mon, grasslands. 

Portulaca umbraticola Kunth ssp. coronata 

Small) Matthews & Ketron [2 coronata 

Small]. Purslane; summer annual. SC: 83717; 

common, uplands. SR: 8286, 8570; uncom- 


— 


n,u 


ands. 
inum aurantiacum Engelm. Flame flower; her 


Tal ‘ {| | 
baceous enn SC: occasional, grass- 
lands. | arasslands 

Talinum paniculatum (Jacq.) Gaertn. Pink baby 


reath 


SC: 8353; common, canyons. 

Primulaceae 

*“Anagallis arvensis De Pimpernel, yeiog ae Pale 
annual. 


Androsace occidentalis Pursh. Rock jasmine; win- 
ter annual. SC: 8827 canyons, up- 
lands. SR:8936, 8950: common, low areas, Sa- 
caton flats. 

Samolus vagans Greene. Water pimpernel; her- 
baceous perennial. SC: 8492; rare, wet soil 
along stream, Fresno Canyon. 


Ranunculaceae 

Anemone tuberosa Rydberg var. tuberosa. Desert 
windflower; herbaceous perennial. SC:8825; 
occasional, rocky slopes. 

Clematis drummondii Torr. & A. Gray. Texas virgin 
bower, barbas de chivito; woody vine. SC: 

403; uncommon, 

Sonoita Creek. 

Delphinium scaposum Greene. Barestem lark- 
spur; herbaceous pais: SC: 9066; locally 
common, uplan 

Delphinium wootonii a berg [D. virescens Nutt. 

JE 


gallery forest along 


a WOON act ]. Larkspur; her- 
| I.SR:9357-rare ae 
Myosurus cupulat S Walco Mouseeilwiaer 


BRIT.ORG/SIDA 22(1) 


annual. SC: 8862, 9010, 9023; abundant, wet 
soil of canyon bottom, north-facing slopes. 
SR: 8934, 9056; abundant, low areas, sacaton 
flats. 
Myosurus minimus L. Mousetail; winter annual. 
SR: 9254, 9350; uncommon, springs. 
Ranunculus hydrocharoides A. Gray. var. 
taeg dasee Buttercup, herbaceous pe- 
ial. SR: 9259; locally co 
aoe macranthus Scheele. Large butter- 
cup;herbaceous perennial. SR: 9485; uncom- 


mmon, springs 


mon 

Thalictrum Pai Engelm.ex A. Gray var.fendleri. 
Meadow rue; herbaceous perennial.SC:8543; 
rare, canyons, north-facing slopes. 


Rhamnaceae 

Ceanothus greggii A. Gray. Buckbrush; shrub. SC: 
9544; rare, Canyons 

Condalia warnockii M. C. Johnston. var. 
kearneyana M. C. Johnston [Condalia 
mexicana Schlecht. sensu Kearney & Peebles 
(1960)]. Mexican bluewood; shrub. SC: 8367; 
occasional, rocky slopes 

Sageretia wrightii S. Watson. Sarl SC: 8671; lo- 
cally common, canyons. 

— obtusifolia (Hook. ex Torr. & A. Gray) A 
Gray var. canescens (A. Gray) M.C. Johnston 
ae lycioides (A. Gray) Weberb.]. Gray 
thorn, abrojo, bachata; shrub. SC: 9494; occa- 
sional, mostly along edge of gallery forest 
along Sonoita Creek. 


osaceae 
*Malus pumilla P. Mill. Apple; tree. SR: 9484, a few 
trees persisting at Sharp Spring near junc- 
tion with Santa Cruz River floodplain 
Potentilla wheeleri S. Watson [P viscidula ayabi 
Cinquefoil; herbaceous perennial. SR: 9059; 
uncommon, north-facing slopes in grass- 
lands, usually found at much higher eleva- 
tions in southeastern Arizona 
Purshia stansburiana (Torr.) Henrickson 
[Cowania mexicana D.Don var. stansburiana 
(Torr) Jespon]. Cliffrose, pofio; large shrub. SC: 
44; occasional, uplands. SR: 8724; uncom- 
mon, grasslands. 
Rubiaceae 
Bouvardia ternifolia (Cav.) Schlecht.[B. glaberrima 
ngelm.]. Shrub. SC: 8435, 9361; rare, north- 
facing slopes, ridgetops. 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 695 


Diodia teres Walt. var. angustata A. Gray. Rough 
buttonweed, poor Joe; summer annual. SC: 
8767. rare, canyons. SR: 8719; uncommon, 
grasslands 

Galium aparine L. Goosegrass bedstraw, cleav- 
ers; annual vine. SC: 9083; common in can- 

ons, below oaks. 

Galium microphyllum A.Gray. Herbaceous peren- 

nial. SC: 9355; uncommon, rock walls. 

Galium ee A.Gray.Great Basin bedstraw; 

annual. SC: 9114, 9275; uncommon, 


canyons. 
Galium Ae aloe perennial. SC: 
02; uncommon, walls. 

Houstonia fled A. cy nce perennial. 
SR: 8 common, north-facing 
slo ssla ae 

bcc breviflorus A. Gray. Summer annual. 
SC: 8 9721; uncommon, ew canyon 
eee 


asslands 


Rutaceae 

Thamnosma texana (A. Gray) Torr. Turpentine 
broom; herbaceous perennial. SC: 9067; un- 
common, uplands 


Salicaceae 
Populus fremontii S. Watson. Fremont cotton- 
wood, dlamo; large tree. SC: commen, gal- 
lery forest al S eek.SR:common, 
springs, Santa Cruz River ek 
Salix bonplandiana Kunth. Bonpland willow, 
uce, sauz; tree. SC: 8651, 8665, 9264; occa- 
sional, gallery forest along Sonoita Creek. 
Salix exilifolia Dorn [Salix taxifolia Kunth]. Yew- 
; small tree. SC: 8669, 9150; 
rare, Sonoita Creek. SR: 9267; uncommon, 
springs, Santa Cruz River bott 
Salix gooddingii Ball. Goodding ee sauce, 
sauz; tree. SC: 8876, 9148; abundant, gallery 
forest along Sonoita Creek. SR: 9429; com- 
mon, springs, Santa Cruz River bottom. 
Salix lasiolepis Benth. var. lasiolepis. Arroyo wil- 
low, sauce; large shrub. SC: 9064; common, 
Sonoita Creek.SR:8949, 91 11,9260, 9262;com- 
mon, springs, Santa Cruz River bottom. 


leaf willow, tardi. 


Sapindaceae 
Dodonaea viscosa (L deed Hopbush, jarilla, tara- 
C:9 canyons 


chiqui;shru 


ged saponaria L. var.drummondii (Ho 
rn.) L.Benson.Western soapberry, amoli ll, 


cirioni, jaboncillo; small tree. SC: 9498; occa- 
er canyons, rocky slopes. SR: 8596; un- 
common, Parker Canyon. 


Scrophulariaceae 

Castilleja exserta (Heller) Chuang & Heckard ssp. 
exserta [Orthocarpus purpurascens Benth,]. 
Owl's clover;hemiparasitic winter annual. SC: 

208; rare, Mesquite grasslands. 

Castilleja integra A. Gray. Indian paintbrush; 
hemiparasitic herbaceous perennial. SR: 
9103, 9251; occasional, grasslands 

Castilleja lanata A sp./anata.\indian paint- 
brush; hemiparasitic herbaceous perennial. 
SC: 8917, 9137, uncommon, canyons 

aoe minor (A. Gray) A. Gray ssp. minor. In- 

aintbrush; hemiparasitic annual. SC: 


ee e, canyon bot : 
Maurandella antirrhiniflora (Humb. & Bonpl. ex 
Willd.) Rothm. [Maurandya antirrhiflora 


Humb.& Bonpl.]. Herbaceous perennial vine. 

SC: 8677: ional, canyons, rock walls. 

Mecardonia procumbens (P. Mill.) Small [M. 
vandellioides (Kunth) Pennell]. Herbaceous 
perennial. SC: 8445, 8468; occasional, canyon 


bottom. 

Mimulus floribundus Lindley. Monkey flower, 
baisémari, herbaceous perennial. SC: 9234; 
rare, Coal Mine Canyon 

Mimulus guttatus DC. Monkey flower, baisomari; 

baceous perennial. SC: 8838; common, ri- 
parian areas. SR: 9096; common, Santa Cruz 


River bottom. 

Mimulus rubellus A.Gray.Red-stemmed mimulus; 
winter annual. SC: 8828; locally common, 
mostly in canyons. SR:9053;occasional, grass- 
lands. 

Nuttallanthus texanus (Scheele) D.A. Sutton 
[Linaria texana Scheele]. Texas toad flax; win- 
ter annual. SC: 8864, 8912; common, rocky 
slopes. SR: 9094; grasslands 

Penstemon barbatus (Cav.) Roth ssp. barbatus. 
Scarlet bugler; herbaceous perennial. SC: 
9407, rare, Fresno Canyon. SR: 9453; rare, oak 
woodlands adjacent to Parker Canyon. 

Penstemon linarioides A. Gray ssp. linarioides. 

ous perennial. SR: 9383; occasional, 


grass a ae 

Penstemon parryi (A. Gray) A. Gray. Herbaceous 
perennial. SC: 9720; occasional, rocky slopes, 
rock walls. 


Penstemon ei (A. Gray) T. J. Howell. 
ngue; herbaceous perennial. SR: 
9525,9568; uncommon, north-facing slopes 
in oak woodlands and grasslands. 
Sairocarpus nuttallianus (Benth. ex A. DC.) D. A. 
Sutton [Antirrhinum nuttallianum Benth,]. 
Winter annual. SC: 9127; rare, rocky slopes. 
Schi alae ee gsi (A. Gray) Pennell. 
annual. SC: 8480; uncommon, 


eae a ee (L.) Sw. Herbaceous peren- 
nial, SC: 9404; rare, rock walls, Fresno Canyon. 
SR: 8734; rare, Parker Canyon. 

Verbascum vi Stokes. Mull bust an- 
nual. SC: 9476; uncommon, along Sonoita 
Creek. SR: 8306; uncommon, springs 

*Veronica anagallis-aquatica L.Water speedwell, 
herbaceous perennial. SC: 9145; occasional, 
riparian areas 

Veronica Sa eate 2 i880. xalapensis (Kunth) 
Pen lly winter annual. SC: 8859 
on 5. sbundant canyons and nid 
drainages. SR:8 es 


Simaroubaceae 
*“Ailanthus altissimus (P. Mill.) Swingle. Tree of 
ven, Copal tree; tree. SC:one large colony 
in the gallery forest of Sonoita Creek. SR:8272; 
Il col | road east of 


¥ dlONnG 


uncommon 


the Santa Cruz River. 


Solanaceae 
Calibrachoa parviflora (Juss.) D'Arcy aad 
parviflora Juss.]. Wild petunia; annual. $ 
9149; uncommon, wet soil along Sono 
Creek. SR:9 


tanks. 
ee ede coniodes ee ex Dunal) Brit- 
ton Small ground cl 
nial. SC: 9296; uncommon, grasslands, SR: 


944]: roadsides 

Chamaesaracha coronopus (Dunal) A.Gray. False 
nightshade; herbaceous perennial. SR: 9367, 
9402; uncommon, roa 

Datura quercifolia Kunth. eon weed toloache; 
herbaceous perennial.SR:8577; uncommon, 
roadsides 

Datura wight Regel [D. meteloides DC]. Sacred 
datura, toloache; herbaceous perennial. SC: 

ccasional, sandy soils adjacent to 

Sonoita Creek. SR:8564; occasional, roadsides, 


BRIT.ORG/SIDA 22(1) 


Lycium berlandieri Dunal var. parviflorum (A.Gray) 
Terracc. Wolfberry; shrub. SC: 8874; uncom- 

on, uplands. 

lycium exsertum A. Gray. Wolfberry; shrub. SC: 
8817, uncommon, uplands. 

Margaranthus solanaceus Schlecht. Netted 
globe berry; summer annual. SR: 8725; un- 
common, oak woodland adjacent to Parker 
Canyon 

*Nicotiana glauca Graham. Tree tobacco, 
corneton, juan loco, palo loco; robust herba- 
ceous perennial. SC: 8693; occasional along 


onoita Creek. 

Nicotiana obtusifolia Martens & Galleotti var. 
obtusifolia [N. trigonophylla Dunal var. 
trigonophylla]. Desert tobacco, tabaquillo; 
herbaceous perennial. SC: 8684; occasional, 


nyon walls 
Physalis acutifolia (Miers) Sandw. [P wrightii A. 
Gray].Wright sual cherry, tomatito; sum- 
mer annual.SR: 8265, 8341: |, grass- 
lands, springs 
Physalis hederifolia A. Gray var. hederifolia. lvy- 
leaved ground cherry, tomatito; herbaceous 
perennial.SR:8588; uncommon, Parker Canyon. 
Physalis longifolia Nutt. Tomatito; herbaceous 
perennial. SR: 8299, 8300, 8706; common, 
springs. 
Physalis pubescens L. var. integrifolia (Dunal) Wa- 
terfall. Hairy ground cherry, tomate, fresadilla; 
DL 


rn dn 


Solanum douglasii Dunal. Nightshade; herba- 
ceous perennial. SC: 8491; rare, Fresno 
Canyon. 

Solanum elaeagnifolium Cav. Silverleaf night- 
shade, trompillo; herbaceous perennial. SC: 
9363; occasional, mesquite grasslands. SR: 
9398; occasional, disturbed areas. 

Solanum rostratum Dunal. Buffalo bur; herba- 
ceous perennial. SC: 9633; uncommon 
mostly along Sonoita Creek. SR: uncommon, 
roadsides 


Sterculiaceae 

Ayenia filiformis S.Watson [A. pusilla L.sensu Kear- 
ney and Peebles (1960)]. Herbaceous peren- 
nial. SC: 8372; occasional, rocky slopes. 

Tamaricaceae 


aaa Moss iin Ledeb. [7. pentandra Pall. 
SETISU KEAINey dla | Peet sles (1 960)] Tamarisk, 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 697 


pino salado; small tree. SR: 9348; rare, Santa 
Cruz River bottom 


Ulmaceae 
Celtis reticulata Torr. Hackberry, palo blanco, 
cumaro; tree. SC common, gallery for- 
ests and canyons. $R:8345; 
f Santa Cruz River floodplain. 
Celtis pallida Torr. Desert hackberry, garambullo; 


— 


edge 
~~ 


rub. SC: uncommon, rocky slopes. 


roe 


rticaceae 
Parietaria bes Hinton var.hespera [P floridana 
Nutt nd Peebles (1960)].Pel- 
litory, Saeuines winter annual. SC: 8830; oc- 
casional, rocky slopes 


Q 


Verbenaceae 
Aloysia wrightii Heller ex Abrams. Wright lippia, 
oreganillo; shrub. SC: 8525; common, rock 
slopes. 
Bouchea prismatica (L.) Kuntze. Summer annual. 
59; uncommon, low areas. 
Glandularia bipinnatifida (Nutt.) Nutt. var. 
bipinnatifida. [Verbena bipinnatifida Nutt]. 
Verbena; herbaceous oe SR: 8261], 
9101;common, grasslan 
Phyla nodiflora (L.) Greene. ee peren- 
nial. SR: 9443; one a near ranch build- 
ings, probably introduc 
Verbena carolina L. Fese-oul perennial. SC: 
uncommon, canyons 
Verbena ae is Desf. orbacealls perennial. SR: 


Small var, 
xylopoda Perry. Hillside vervain; herbaceous 
perennial. SC: 8478, 8739; occasional, mostly 
on shaded slopes. 


Necheeae 
alifornicum Nutt. Desert mistle- 

toe; enaicl herbaceous epiphyte. SC: 
8849, occasional on Prosopis, Condalia. 

Phoradendron capitellatum Torr. ex Trel. [P. 
bolleanum (Seem.) Eichler var. capitellatum 
(Torr. ex Trel.) Kearney & Peebles]. Mistletoe; 

| us ee SC:8685, 


hemiparastic I 


er coryae Trel, ene hemipara- 
sitic herbaceous epiphyte. SC: 8868; occa- 
sional on Quer blongifolia. SR: : 
casional, on Ouen rCUS SPP. 

Phoradendron tomentosum (DC.) Engelm. ex A. 


Gray [Pflavescens (Pursh) Nutt. var. pubescens 
Engelm.]. Mistletoe, inierto; hemiparasitic 
herbaceous epiphyte. SC: 9018; uncommon 
on Fraxinus 


Vitaceae 

Vitis arizonica Bnet) sn grape, parra del 
monte; woody vine. 417;occasional,can- 
yons. SR: 9395; nae common, Parker 
Canyon. 


Zygophyllaceae 

Kallstroemia grandiflora Torr.ex A.Gray. Mexican 
poppy, summer poppy, baiburin, mal de ojo; 
summer annual. SC: 8361; occasional, can- 
yons and slopes. 

Kallstroemia parviflora Norton. Caltrop; summer 
annual.SR:8339 
Parker Canyon. 

*Tribulus terrestris L. Caltrop; summer annual. SR: 
9591; uncommon, disturbed sites. 


8713;:occasional, grasslands 


MAGNOLIOPHYTA: LILIOPSIDA 


Agavaceae 

Agave palmeri Engelm.Palmer agave, lechuguilla; 
large leaf succulent. SC: occasional, canyons 
and rocky slopes. SR:uncommon, grasslands. 

Agave parryi Engelm. var. parryi. Century plant; 
large leaf succulent. SR: uncommon, grass- 
lands. 

Agave schottii Engelm. var. schottii. Shindagger. 

amolillo ll leaf succulent. SC: 9413; 
bundant on uplands 

Yucca elata Engelm. Soap tree yucca, palmilla; 
caulescent leaf succulent. SC: 9408; uncom- 
mon, canyons. SR: rare, a single individual 
observed in grassland west of ranch house. 

Yucca schottii Engelm. Mountain yucca; 
caulescent leaf succulent. SC: uncommon, 
north- facing slope, Coal Mine Canyon. SR: 
rare, a single individual observed with oaks 
east of Santa Cruz 

Yucca oe ee Banana yucca, datil: 
acaulescent or short-caulescent leaf succu- 
lent. SC: abundant, uplands. Kartesz (1999) 
treats this as a variety of Yucca baccata; the 
latter taxon has strongly glaucous foliage, 
thick leaf fibers, and tends to form dense 
colonies of acaulescent rosettes. The plants 
of southern Arizona have green foliage, thin- 
ner fibers, and are often caulescent 


amol:sma 


698 


Alismataceae 

Sagittaria se Engelm. ex J. G. Sm. Arrow- 
head; aquatic perennial. SC: 6488, 
J.Kaiser 1 ber ae collected four times from 
a Cattle tank adjacent to Santa Cruz River, 
formerly plentiful’; last collected in 1991, 


Bromeliaceae 

Tillandsia recurvata (L.) L. Ball moss, gallitos; epi- 
La leaf Succulent, SC: 8678; locally abun- 

niperus coahuilensis 


in Narrow Canyons. 


Commelinace 
Commelina di itl ifolia Delile. Day flower, 
tale de Maria; herbaceous perennial. SC: 
ons and rocky slopes. 
n, grassland 
Coramnelien erecta L. var. angustifolia (Michx.) 
Fern. Day flower, yerba del pollo; herbaceous 
perennial.SC:8355;uncommon,canyons and 
rocky slopes. SR: 9622; uncommon, Parker 


SR 8314 


Canyon. 
Tradescantia pinetorum Greene. Spiderwort; her- 
aceous perennial. SR: 9565; rare, grasslands. 


Cyperaceae 

Carex praegracilis W. Boott. Sedge, herbaceous 
i re ae SR: 9248, 253, ‘ona 9463; abun- 

|\ y collections are 

atc to eran om C.chihuahuensis 


= 


ackenzie. 
oped Liebamann [C. wrightii aaa 
edge; herbaceous perennial. SC: 8 
common,north-facing slopes below oa 
ee esculentus L. eed hutseelge) herba- 
ceous perennial. SC: 9 an- 
yons. SR: 9593, 9598, 9611 7 occasional, grass- 
lands, roadsides. 

Cyperus none Michx. [C. albomarginatus 
Mart. & Schrad.]. Flat sedge; herbaceous pe- 
rennial. SC: 8467, 8539, 9647; common along 

ream in Fresno Canyon. 

Cyperus mutisii (Kunth) Griseb. Flat sedge; her- 
baceous perennial. SC: 9708; rare, Fresno 


Canyon, 
ne niger Ruiz & Pavon. Flag sedge; herba- 
ous perennial. SC: 8776; uncommon, 
a Canyon. SR:8324,8789, 9599;common, 
nta Cruz River bottom 
ope odoratus L. Flat sedge; herbaceous pe- 
rennial.SC:8495, 9668, 9684; common in wet 


BRIT.ORG/SIDA 22(1) 


soil ae Sonoita Creek. SR: 9668; common, 
z River bottom. 

Cyperus pal iin or (Kukenthal) G. Tucker [C. 
flavus (Vahl) Nees, in part]. Flat na her- 
baceous perennial. SC:8438; rare, below 

cane sphaerolepis Boeckl.[C. rusbyi en i 

ge; herbaceous perennial. SR:8508; occa- 
he oak sav 

Cyperus SqUBHOSUS ai [C. aristatus Rottb.]. Flat 

sedge; annual. SC: 9497, 9640; uncommon, 
moist af of canyon bottom 

Eleocharis bella (Piper) Svenson. Spike rush;sum- 

ual. SR: 9575 |, Santa Cruz 
River bottom. 

Eleocharis montevidensis Kunth. Spike rush; her- 

; -41 SC-939()-a} 


along 


streams. SR: 9437; abundant, springs, Santa 
Cruz River bottom 
Eleocharis palustris (L.) Roemer & J.A.Schultes [. 
macrostachya Britton sensu Kearney & 
Peebles (1960), in part]. Spike rush, herba- 
ceous perennial. SC: 9293: occasional, mostly 
in canyons. nr aca all 
Cruz River bot 
Fimbristylis annua a Roemer & J.A. Schultes 
[F. baldwiniana (Schultes) Torr]. ara an- 
nual. SC:8539; uncommon, canyon bot 
Lipocarpa micrantha (Vahl) G. Tucker heee 
carpha micrantha (Vahl) ie Summer an- 
nual. SC: 9675; n botto 
sae ore acutus (Muhl. ex oa) A. ‘é 
ssp. occidentalis (S.Watson) $.G.Sm. 
ae acutus Muhl.]. Bullrush; herbaceous 
perennial. SR: 9461; uncommon, springs. 
Schoenoplectus americanus (Pers.) Volk.ex Schinz 
&R. a istaabe americanus assets Bullrush; 


CPOUS 
p 


— 


Ut] MT IOrd, 


Santa Cruz River bottom. 


lridaceae 

Sisyrinchium demissum Greene. Blue eye grass; 
herbaceous perennial. SR: 9376; common, 
springs 


JUNCACEAE 
Juncus bufonius L. Toad rush; annual. SC: 97.73, 
mmon, moist soils of canyon bottom. 
Juncus interior i eeaealereaciae a perennial. 
SC: 9236, 9294 ms. 
Juncus mexicanus Willd. ex J A & J.H. Schultes. 
Rush; herbaceous perennial. SR: 9325, 9388, 


gq 
=) 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATU 


AREA AND SAN RAFAEL STATE PARK 699 


9430, 9433; abundant, springs, Santa Cruz 
River bot 

Juncus saximontanus A. Nelson. Rush; herba- 
ceous perennial.SC:8493, 9210;uncommon, 
along streams. SR: 9315, 9438, 9464, 9487. 

_ springs, Santa Cruz River bottom. 

Juncus torreyi ae Rush; herbaceous peren- 
nial. SR: 8284, 9435; common, springs, Santa 
Cruz River bottom. 


commo 


{ {| €] 


eae 

Lemna gibba L.Windbags, duck 
ing aquatic. SC: rare, Fresno Canyon. SR:8373; 
occasional, springs. Usually found as a minor 
component in mixed colonies with L. minor. 

Lemna minor L. Water lentil, duckweed: small 
floating aquatic. SC: 8499; locally abundant 
in pools of canyons. SR: 8372; common, 
springs 

Liliaceae 

Allium kunthii G. Don. Wild onion; herbaceous 
perennial. SC: 9644; rare, Coal Mine Canyon. 

Calochortus ambiguus (M.E. Jones) Ownbey. 
Mari lily;hert ial. SC: 9282. 


locally abundant, Fresno Canyon. 
Calochortus kennedyi Porter var. kennedy. so 
ariposa; herbaceous perennial. SC: 9724; 
occasional, uplands. SR: 9241; locally com- 
mon, grasslands. 

Dichelostemma capitatum (Benth.) Wood ssp. 
pauciflorum (Torr.) G. Keator [D. pulchellum 
(Salisb.) Heller].Bluedicks; herbaceous peren- 
nial. SC: 8929; common, uplands. SR: 9106; 

grasslan 

Pee as pee U. 7 & H.H.Schultes) Cruden 
[Anthericum torreyi Baker]. Crag lily; herba- 


eous perennial SR: 95278 


grasslands. 

Milla biflora Cav. Mexican Star, herbaceous pe- 
rennial. SC: rare, included based on a photo 
by R. Gagnon. SR: 9631; rare, grasslands 

Nothoscordum texanum M.E. Jones. Herbaceous 
perennial.SC:8440, 9235; uncommon,mostly 
in seeps in Coal Mine Canyon. 


Nolinaceae 

Dasylirion wheeleri S. Watson. Desert spoon, so- 
tol; leaf rosette shrub. SC: common, rocky 
slopes. 

Nolina microcarpa S. Watson. Beargrass, 
sacahuista, palmilla; leaf rosette shrub. SC: 


common, rocky slopes, grasslands. SR: locally 
abundant, grasslands and oak wooclands. 


Poaceae 
Agrostis exarata Trinius. Spike bent; herbaceous 
perennial. SR: 9439; Santa Cruz 


Alopecurus carolinianus Walt. Fox tail; summer 
annual. SR: 9255; uncommon, springs. 

Aristida adscencionis L. Six weeks three-awn; 
summer annual. SC: 8426, 9229: common, 
uplands. SR: 9588; occasional, grasslands, 

dsides. 

Aristida divaricata Humb,& Bonpl.ex Willd. Three- 
awn; herbaceous perennial. SR: 9698; occa- 
sional, grasslands 

Aristida purpurea Nutt. var. fendleriana (Steud.) 
Vasey [A. fendleriana Steud.]. Purple three- 
awn; herbaceous perennial. SR: 9374; occa- 
sional, grasslands 

Aristida purpurea Nutt. var. neallyi (Vasey) Allred 
a glauca ise ia SC: 9321, 9543; u 

n, uplan 

Pee purpurea ea var. wrightii (Nash) Allred 

A. wrightii Nash]. Purple three-awn; herba- 

ceous perennial. SC: 8573; occasional, grass- 

lands. SR: 9605; common, gra asslands 

ristida ternipes Cav. var.gentilis (Henr.) A lred [A. 

mulosa Henr]J. Spider grass; herbaceous 
perennial SC: 8428; common, canyons. SR: 
grasslands 


= 


co 


Asti as Cav. var. ternipes. SC: 8505, 8520 
ceous perennial; Common, canyons. 
mt fatua L.Wild oats; winter annual. SC: 9238; 
rare, canyons. SR: 9264; uncommon, dis- 
turbed areas. 
*Avena sativa L. Oats; winter annual. SR: 9752; 
incommon, disturbed a 
birch barbinodis a Herter [Andro- 
rbinodis Lag.J. Cane beardstem; 
feat perennial. SC: 8430; abundant, 
uplands. SR: 8329; abundant, grasslands. 
Bothriochloa laguroides (DC.) Herter ssp. 
torreyana (Steud.) Allred & Gould [Andro- 
pogon ISO Swartz]. alee 
rp Spring 


perennial.S 


drainage. 
eee aristidoides (Kunth) Griseb. var. 
aristidoides. Six-weeks needle grama; sum- 
mer annual. SC: 8434; occasional, canyons 
and uplands. SR: 8572; common, grasslands 


700 


Bouteloua chondrosioides (Kunth) Benth. ex S. 
Watson. Spruce-top grama; herbaceous ES 
ee SC:8433,8502;common,uplands.SR 
8708 grasslands 


— curti inenctale (Michx.) Torr. Side oats 
erbaceous perennial. SC:8402; abun- 
ee canyons and uplands. SR: 8550; abun- 


dant grasslands woodlands 


sasaueson a ae selects peren- 
nial.S and 


adjacent to 


BRIT.ORG/SIDA 22(1) 


areas. SR: 8549; | grasslands, road- 


sides. 
Cottea pappophoroides Kunth. Cotta grass; her- 
baceous perennial. SC: 9679; rare, Fresno 


Canyon. 
*Cynodon dactylon (L.) Pers.Bermuda grass; her- 
aceous perennial. SC: 9284; occasional, 
mostly along streams. SR: 9563; ss aahdiad 
fale Santa Cruz River botto 
pulchella (Kunth) Willd. ex caviar 


Parker ee 
Bouteloua ene ou (Torr.) Torr. Black grama;her- 
s perennial. SC: 9637; uncommon, 
tee aa SR: 9604; sslands 
Bouteloua gracilis (Willd. ex it Lag. ex 
Griffiths. Blue ist eae perennial. 
grasslands 


~~ 


SR: 9 
Bouteloua hirsuta Lag. var. aa Hairy grama; 
NELDacSOUs Perennial SC: 8429; common, 
uplands grasslands 
Bouteloua railcar (E. Fourn.) Griffiths. Herba- 
eous perennial. SC: 8761; occasional, up- 
lands. SR: 9626; uncommon, oak woodland 
adjacent to Parker Can 
Bouteloua repens (Kunth) cation: & Merr. [B. 
filiformis (Fourn.) Griffiths]. Slender grama; 
erbaceous perennial. SC: 8427; common, 
uplands. 
Bouteloua rothrockii Vasey. Rothrock grama;her- 
ceous perennial. SC: 8528A; occasional, 
mesquite grasslands. SR: 9601; occasional, 
low areas, sacaton flats. 
Bromus anomalus Rupr. ex E.Fourn. Brome; her- 
eous perennial. SR: 9696; rare, Parker 


Canyon. 
Bromus carinatus Hook. & Arn. California brome; 
herbaceous perennial. SC: 9215, 9267; com- 
mon, gallery ies along Sonoita Creek. 
*Bromus catharticus Vahl.Brome; annual. SR: 9052 
9158, 7 commen Sn Cruz River flood- 
plain, disturbed are 
*Bromus japonicus =n ex Murr. Japanese 
bro rama: annual. SR: 9389; common, 
Sharp Spring drain 
pence, spinifex a c nay orus Benth. 
Sandbur;summer annual.SC:8667;common, 
mostly in su ae ieamaa areas. SR:85 76; 
mmon, grassla 
Chloris virgata her fingergrass; sum- 
mer eae SC: 8398B; its, disturbed 


Swal L 


pee ‘pulchellus (Kunth) Hitchc.]. Fluff 
grass; herbaceous perennial. SC: 9547, 9648; 
locally common, uplands 

Digitaria californica (Benth.) Henr. [Trichachne 
californica (Benth.) Chase]. Arizona 
cottontop; herbaceous perennial. SC: 8432; 
uncommon, uplands. SR: 9609; uncommon, 
grasslands. 

Digitaria cognata (J.A. Schultes) Pilger var. 
pubiflora Vasey ex L.H. Dewey [Leptoloma 
cognatum (Schultes) Chase]. Fall witch grass; 
teease| “41 SC-9028 


ae 


Coal Mine Canyon. 

“Digitaria sanguinalis (L.) Scop. Common c 
grass; summer annual. SC: 8660; ator 
wet areas. SR: 9589; occasional, disturbed 


areas. 
“Echinochloa colana (L.) Link. Jungle rice; sum- 
mer annual. SC: 8537; ional, canyons 


*Echinochloa crus-galli (L.) P. Beauv. Barnyard 
rass; summer annual. SC: 8452, 8487; occa- 
sional, along streams. SR: 8269; common, 
Santa a River bottom, abundant around 
cattle 

Elionurus ae tibial s Hack. Herbaceous peren- 
nial. SC: 97108; rare, shaded slopes 

Elymus canadensis L.Wild rye; herbaeous peren- 
nial. SR: 94 Santa Cruz River bot- 
tom, springs. 

Elymus elymoides (Raf.) Swezey [Sitanion hystrix 
(Nutt.) J.G. Smith]. Bottlebrush squirrel tail; 
herbaceous Lear SC: 91] 8; Secsleiiel 

R: 9263; |, Sharp 


canyons, slopes 
Spring drainage. 
Enneapogon desvauxii Desv. ex Beauv. Spike 
pappusarass;summer annual. SC:8474, 8482; 
uncommon, rocky slopes. SR: 8777; uncom- 


n, grasslands. 
eet ae (All.) Vign.ex Janchen. Stink 
grass; annual. SC: 8450; mostly in heavily 


MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL 


AREA AND SAN RAFAEI STATE PARK 701 


Cp.ea74 


grazed areas. SR: 82 


|, disturbed 


areas. 
*Eragrostis curvula (Schrad.) Nees var. conferta 


nial. SC: 9277; 
eee intermedia Hitchc. Plains lovegrass; 
er ePSTeniich SC: 8439; common, 

me a 8548 dant, grasslands 
ks lehmanniana Nees. Lehmann 
vegrass; herbaceous perennial. SC: 9499; 
Bi mostly oe saddles. SR: 
9483: locally co 
Eragrostis pectinacea (Michx.) Nees ex Steud. var. 
pectinacea [E. diffusa Buckley]. Summer an- 
ual. SC: 8530, 9495: occasional, canyons. SR: 
9612; occasional, grasslands, ene 

Eriochloa acuminata (Presl) Kunth var.acuminata 
(Vasey) R.B. Shaw [E. gracilis (Fourn.) Hitche. 
var. gracilis (Vasey) Hitchc.]. Cupgrass; sum- 
mer annual. SC: 8375; common, gallery for- 

ests, Canyons. 
Eriochloa acuminata (Presl) Kunth var. minor 
(Vasey) R.B. Shaw [E. gracilis (Fourn.) Hitch. 
var.minor (Vasey) pcue Cupgrass;summer 


canyons 


mmon, grasslan 


LO) 
lox 
oO 2 


se 


ual. SR: QI Hl + | 
OL 


ann 

ne arundinacea Schreber. [F. elatior L.]. 
Meadow fescue; herbaceous perennial. SR: 
9299, 9314, 9352; abundant, springs 

Heteropogon contortus (L.) Beauv. ex Roemer & 
J.A.Schultes. Tangle head; herbaceous peren- 
nial. SC: 8770; rocky slopes 

Heteropogon melanocarpus (EIl.) Ell. ex Benth. 
Sweet tangle head;summer annual. SC:8396; 
common, rocky slopes. SR: 8702; occasional, 
oak savannas. 

Hilaria belangeri (Steud.) Nash var. belangeri. 
Curly mesquite grass; herbaceous perennial. 
SC: 8816; locally abundant, grasslands. SR: 

9581; occasional, grasslands. 

Hordeum jubatum L.ssp.jubatum. Foxtail;herba- 
ceous Beicuael SR: 9308B, 9349; common, 
springs, low areas 

*Hordeum murinum L. ssp. leporinum (Link) 
Arcang. Wild barley; winter annual. SR: 9049; 

asional, margins of cattle tanks 

Hordeum eid Nutt. Little barley; annual. SR: 

8A; uncommon, margins of cattle tanks. 

Leersia oryzoides (L.) Sw. Rice cutgrass; herba- 

ceous perennial. SR:8783, 8786; uncommon, 
Sharp Spring. 


Leptochloa dubia (Kunth) Nees.Green sprangle- 
top; herbaceous perennial. SC:8421;uncom- 
mon, rocky slopes. SR:8707; occasional, grass- 
lands. 

ec oa fusca (L.) Kunth ssp. fascicularis (Lam.) 

w [L. fascicularis (Lam.) A. Gray var. 
aaa Beaded sprangletop; summer 
i aoe 
disturbed are 

Leptochloa ie (Retz) Ohwi ssp. lies 
(Steudl.) N. Snow [L. filiformis Lam.].Sum 
annual. SC: 8529; common, canyons. 

Leptochloa viscida (Scribn.) Beal.Summer annual. 
SC: 9496; occasional, mostly in canyons. SR: 

F mmon, washes. 

Lycurus setosus (Nutt.) C. G. Reeder [L. phleoides 
Kunth sensu Kearney and Peebles (1960), in 
part]. Wolf-tail; herbaceous perennial. SC: 
8425;common, uplands. SR: 9570; abundant, 


— 


grasslands. 

*Melinis repens (Willd.) Zizka [Rhynchelytrum 
roseum (Nees) Stapf & C. E. Hubbards ex 
Bews]. Natal grass; herbaceous perennial. SC: 

813:rare, canyons. SR: 9736; rare, Parker Can- 
yon. 


NArihl 


ica Scribn. Herbaceous pe- 

enna SC: 8760, 9078; locally abundant, up- 
lands. SR: 9658; uncommon, grassland 

Muhlenbergia oe (Nees & ee ex 
Trinius) Parodi. Scratchgrass; herbaceous 
perennial. SR: 9656; locally abundant, Sharp 
Spring drainage. 

Muhlenbergia emersleyi Vasey. Bullgrass; herba- 
ceous perennial. SC: 8676, 8762; occasional, 
mostly north-facing slopes. 

a fragilis Swallen. Summer annual. 

; asional, canyons and rocky 
oe SR:8730, 9725; uncommon, oak wood- 
land adjacent to Parker Can 

a ate rigens (Benth. oe Deer grass; 
herbaceous ane SC: 8764; abundant, 

m. SR: 9686; occasional, drain- 


canyon b 


ages. 

Muhlenbergia tenuifolia (Kunth) Trinius [M. 
monticola Buckley]. Herbaceous perennial. 
SC: 9642A, 9676, 9705; occasional, rock walls. 

Panicum bulbosum Kunth. Bulb panicum;herba- 
ceous perennial. SC:8378,8507; uncommon, 
canyons. SR: 9628; uncommon, oak wood- 
land adjacent to Parker Canyon. 


Panicum hallii Vasey var. hallii. Herbaceous pe- 
ial 


SR: rare, grasslands 


Sacon hirticaule J. Presl. Summer annual. SC: 
8391, 8395, 8449; common, canyons and 
slopes. SR: 8285; common, grasslands. 

Panicum obtusum Kunth. Vine mesquite; herba- 
ceous perennial. SC: 9549; uncommon, 
washes. SR: 9578; occasional, grasslands, 


springs. 
“Paspalum dilatatum Poir. Dallis grass; herba- 
ceous Sal SC:8664; streams 
SR: 832 |, Sharp Spring. 
Paspalum - stichum L. Knotgrass; herbaceous 
perennial. SR: 8788; locally common, Santa 
Cruz River bottom 
Paspalum setaceum Michx 
ceous perennial. SR: 8 


.[Pstramineum Nash]. 
05, 9595: un- 


common, Oak 5 S. 
Phalaris caroliniana Walt. Canary grass; annual. 
SR: 9309, 9428: occasional, low areas. 
*Phalaris minor Retzius. Littleseed cama grass; 
ual. SC: 9204; /canyon 
Poa bigeloviiVasey & Seti: Bigelow's pie ae 
winter annual. SC canyons 
rocky slopes. 
*Poa pratensis L. ssp. pratensis. Kentucky blue- 
grass; herbaceous perennial. SR: 9256; abun- 
ant, Sharp and Heron springs drainages. 
*Polypogon monspeliensis (L.) Desf. Rabbitfoot 
grass; annual. SC: 9147; common along 
streams. SR: 9099, 9372; anta Cruz 


River bottom. 
*Polypogon viridis (Gouan) Breistr. [Agrostis 
semiverticillata (Forsk.) C. Christens.]. Water 
nt; herbaceous oe aia 9331, 9356; 


Schizachyrium cirratum (Hack.) Wooton & 
aes reo cirratus Hack.]. Texas 
bluestem; herbaceous oo SR: 9668; 
decal aca peer 

ae Altson var. 

ae Mees) | Hatha [Andropogon 

hirtiflorum (Nees) Kunth]. Herbaceous peren- 
nial. SC: 8545, 9681, 9710A; occasional, can- 
yons, shaded slopes. SR: 8785; occasional, 


nor 
Setaria grisebachii E. Fourn. Grisebach 
bristlegrass; summer annual. SC: 8538; occa- 


BRIT.ORG/SIDA 22(1) 


sional, north- — SR: 8792, 9620A; 

oc nal, Parker C 

Setaria irucouie ccibn He og K. Schum. [S. 
macrostachya Kunth sensu Kearney and 
Peebles (1960), in part]. sop bristle grass; 
herbaceous a ape SC:8441,8528B; occa- 

e grasslan 

ee ee (L.) Pers. eee grass;her- 
baceous perennial. SC: 9652; occasional, 
mostly along Sonoita Creek. SR: 9561; locally 
gene roadsides, springs, Santa Cruz 
River floodplain 

Pre abuse (Michx.) Scribn. Prairie 
wedge grass; herbaceous perennial. SR: 9460; 
uncommon spr in 

Sporobolus contractus aeiche Spike dropseed; 
herbaceous perennial. SC: 9635; occasional, 
along Sonoita Creek. SR: 9624; rare, Parker 


Canyon. 

Sporobolus cryptandrus (Torr.) A. Gray. Sand 
dropseed; herbaceous perennial. SR: 9592, 
9611B,9695;uncommon, grasslands, washes. 

Sporobolus wrightii Munro ex Scribn. Sacaton. 
Herbaceous perennial. SC: rare, canyons. SR: 

2,abundant on Santa Cruz River flood- 


plain. 
Trachypogon spicatus (L.) Kuntze [T. secundus (J. 
Presl) Scribn.]. Srinkis: awn; dali S pe- 


rennial.SC:8536,87 arte fac- 
ing slopes. SR: 9608, 9664; common, grass- 
lands. 


Urochloa arizonica (Scribn. & Merr.) O. Morrone 
& F. Zuloaga [Panicum arizonicum Scribn. & 
Merr]. Summer annual. SC: oe can- 

S. 4, occasional, low 

Vulpia aibiosaeiue (Nutt.) Gai var. ee 
(Scribn. ex Beal) Lonard & Gould [Festuca 
eae Piper]. Winter annual. SC: 8870; oc- 

asional, rocky slope 

Vulpia pene a (Walt.) a var. hirtella (Piper) 

He Aa octoflora Walt. var. hirtella 
pict Six-w ue; winter annual. SC: 
8923: ene rocky slopes. SR: 9095; oc- 
casional, grasslands. 

Vulpia octoflora (Walt.) Rydb. var. octoflora. Six- 
weeks fescue; winter annual. SC: 908]: com- 
mon, grasslands, sandy soils along Sonoita 
Creek. 


Pontederiaceae 
Heteranthera limosa (Sw.) Willd. Mud plantain; 


MCLAUGHLIN, FLORA OF 


emergent aquatic perennial. SR: 6489: last 
collected from pond adjacent to Santa Cruz 
River in 1991. 

Potamogetonaceae 

Potamogeton foliosus Raf. Pondweed; sub- 
merged aquatic perennial. SR: 9462, 9465; 


AREA AND SAN RAFAEL STATE PARK 703 


tion seen along Sonoita Creek. SR: 9574: 
uncommon, Santa Cruz River bottom. 


Zannichelliaceae 

Zannichellia palustris L. Horned pondweed,com- 
mon poolmat; submerged herbaceous 
aquatic. SC: 9332; rare, found once in pool 


uncommon, Sharp and Heron springs. along Sonoita Creek. SR: 9245: common, 


Typhaceae springs, Santa Cruz River. 
Typha domingensis Pers. Cattail; emergent peren- 
nial aquatic. SC: rare, a single small popula- 


ACKNOWLEDGMENTS 


I want to extend particular thanks to Betsy Lewis for her capable assistance 
with the field work, often under very difficult conditions. Matt Chew, formerly 
with Arizona State Parks, helped in initiating this study. Rick Gagnon, Lee 
Eseman, Carol Bercich, Dave Pawlik, and Steven Haas of State Parks provided 
extensive logistic support. Access to Sonoita Creek State Natural Area through 
private property was granted by Dave Blouin, Morningstar Ranch, and by Javier 
Badillo. Several individuals helped with the field work at various times; | par- 
ticularly want to acknowledge the assistance of George Montgomery, Rick 
Gagnon, Janice Bowers, Anne Thorne, Jim Jordan, Mills Tandy, Bob Zahner, and 
Glenda Zahner. Erika Geiger drafted the maps of the study areas. Richard S. 
Felger and Dale S. Turner provided very thorough and thoughtful reviews. 


REFERENCES 


Benson, L. 1969. The cacti of Arizona. 3rd ed. University of Arizona Press, Tucson. 

Bock, J.H.and C.E. Bock. 2000. The view from Bald Hill: Thirty years in an Arizona grassland. 
University of California Press, Berkeley. 

Brown, D.E. 1994. Biotic communities: southwestern United States and northwestern 
Mexico. University of Utah Press, Salt Lake City. 

Burcess, T.L. 1995. Desert grassland, mixed shrub savanna, shrub steppe, or semidesert 
scrub? The dilemma of coexisting growth forms. |n:M.P. McClaran and T.R.Van Devender, 
eds. The desert grassland. The University of Arizona Press, Tucson, Arizona. Pp. 31-67. 

Haotey, D.and T.E. SHerioan. 1995. Land use history of the San Rafael Valley, Arizona (1540- 
1960). USDA Forest Service General Technical Report RM-GTR-269. Fort Collins, 
Colorado. 

Kartesz, J.T.1999.A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. 1st ed. In:J.J. Kartesz and C.A. 
Meacham. Synthesis of the North American Flora, Version 1.0.North Carolina Botanical 
Garden, Chapel Hill. 

Kearney, TH. and R.H. Peestes. 1960. Arizona flora. 2nd ed. The University of California Press, 
Berkeley. 


704 BRIT.ORG/SIDA 22(1) 
McLAUGHLIN, S.P. 1989. Natural floristic areas of the western United States. J. Biogeogr. 16: 


McLAauGHUuwN, S.P. 1992. Are floristic areas hierarchically arranged? J. Biogeogr. 19:21-32. 

Myrick, D.F. 1975. Railroads of Arizona. Vol. 1. The southern roads. Howell-North Books, 
Berkeley. 

Parritt, B.D. and A.C. Gisson. 2003. Cactaceae Jussieu. In: Flora of North America Editorial 
Committee, eds., Flora of North America. Volume 4. Magnoliophyta: Caryophyllidae, 
part 1.Oxford University Press, New York. Pp. 92-257. 

RicHarDsON, M.L., S.D. Clemmons, and J.C. Wacker. 1979. Soil survey of Santa Cruz and parts of 
Cochise and Pima counties, Arizona. USDA Soil Conservation Service and Forest Ser- 
vice, Washington, DC. 

Rotuins, R.C. 1993. The Cruciferae of Continental North America: systematics of the mus- 
tard family from the arctic to Panama. Stanford University Press, Stanford, California. 
Setters, W.D., RH. Hitt, and M. Sanperson-Rae. 1985. Arizona climate: the first hundred years. 

University of Arizona, Tucson. 

Wuite, S.S. 1948. The vegetation and flora of the Rio de Bavispe in northeastern Sonora, 

Mexico. Lloydia 11:229-302. 


ANNOTATED CHECKLIST OF THE VASCULAR FLORA 
OF THE BIG SANDY CREEK UNIT, 
BIG THICKET NATIONAL PRESERVE, TEXAS 


Barbara R. MacRoberts and 


Larry E.Brown Michael H. MacRoberts 
Houston Community College Bog Research 
1300 Holman mbia 
Houston, Texas 77004, U.S.A. Shreveport, Louisiana 71104, U.S.A. 
and an 
Herbarium, Spring Branch Science Center Herbarium, Museum of Life Sciences 
8856 Westview Drive Louisiana State University in Shreveport 
Houston, Texas 77055, U.S.A. Shreveport, Louisiana 71115, U.S.A. 


Paul A.Harcombe, 


Warren W. oe |. Sandra Elsik Stanley D. Jones 
Department of Ecology volutionary Biology Herbarium, Botanical ae Center 
Rice Universi’ PO. Box 6 
Houston, Texas 77005, U.S.A. Bryan, Texas sr U.S.A. 
ABSTRACT 


An annotated, vouchered checklist is provided of the vascular plant taxa of the 5,806 hectare Big 
Sandy Creek Unit, Big Thicket National Preserve, Polk County, southeastern Texas. Documented for 
the unit are 693 total taxa with 45 of them exotic 


RESUMEN 


AAs a poe | ] Bead Pane oe Oa Ven | 


Se ofrece un listado con anotaciones, y 
de Big Sandy Creek Unit, Big Thicket National oe Polk ore Sureste de Texas. Se documentan 
para la unidad un total de 693 de los que 45 


INTRODUCTION 


This is the third installment of an inventory, based upon available voucher speci- 
mens, of the vascular flora of the Big Thicket National Preserve, southeastern 
Texas. In the two previous papers we provided an annotated checklist of the 
Hickory Creek and the Turkey Creek units (MacRoberts et al. 2002b; Brown et 
al. 2005.). In this paper we do the same for the Big Sandy Creek Unit. 


SITE AND METHODS 


The Big Thicket is located mostly within the longleaf pine region of the West 
Gulf Coastal Plain in southeastern Texas (Parks & Cory 1936; McLeod 1971; 
Harcombe & Marks 1979; Marks & Harcombe 1981; Harcombe et al. 1993). The 
Big Thicket National Preserve consists of 12 units scattered over seven Texas 


SIDA 22(1): 705 — 723. 2006 


706 BRIT.ORG/SIDA 22(1) 


counties. The units range from 222 to 10,100 hectares and total about 34,000 
hectares (Peacock 1994). The Big Sandy Creek Unit consists of 5,806 hectares 
and is located in eastern Polk County (Fig. 1). It is about 14.5 km from north to 
south and is about 9.7 km at its widest east-west point. In elevation it ranges 
from about 53 to 94 meters. The Big Sandy Creek Unit was chosen for inclusion 
in the Big Thicket National Preserve because it is diverse, has a wide range of 
plant communities, is a stream corridor unit, and has “hilly” topography ab- 
sent from most of the other units (Ajilvsgi 1979; Watson 1979; Harcombe & 
Marks 1981). It is dominated by upper slope pine oak forest, mid slope oak pine 
forest, and lower slope hardwood pine forest. Also present are upland pine for- 
est, floodplain hardwood forest, floodplain hardwood pine forest, baygall, cy- 
press-tupelo swamp, and sandhill pine forest (Harcombe & Marks 1979). It has 
one pitcher plant bog that is badly degraded , no wetland pine savanna, no prai- 
ries, and no xeric sandylands although some of the upper slope pine oak forest 
and sandhill pine forest approach xeric sandylands conditions. A map show- 
ing the distribution of plant community types is in Harcombe and Marks (1979). 
Fire suppression over the past 30 years has resulted in extensive midstory de- 
velopment and shading out of herbaceous vegetation. Details regarding soils, 
topography, and climate can be found in Deshotels (1978), Watson (1979), 
Harcombe and Marks (1979), and Marks and Harcombe (1981). 

Like all units of the Big Thicket National Preserve, the Big Sandy Creek 
Unit had not previously been the subject of a detailed floristic inventory al- 
though it is one of the best documented units (Watson 1982). 

The MacRoberts collected 446 specimens, representing 348 taxa, during 
Ll field days beginning 3 April 2003 and ending 2 May 2004. Larry Brown col- 
lected 644 specimens , representing 435 taxa, during 12 field days beginning 4 
April 2003 and ending 27 September 2005. Stanley Jones collected 252 speci- 
mens in 2003. Geraldine Watson collected 255 specimens over several years in 
the late 1970s and early 1980s. Other collectors gathered about 84 specimens. 
We searched the Texas A & M University (TAMU) herbarium consortium data- 
base for additional specimens but no new taxa were found there. 

Since our purpose is to produce a list of documented taxa known to occur 
on the Big Sandy Creek Unit, a vouchered specimen was considered to be the 
only evidence acceptable for inclusion in the list. However, two species, Phlox 
nivalis subsp. texensis a federally listed species, and Utricularia inflata were 
photographically documented and not collected. In all, about 1,650 herbarium 
specimens form the data for this report. All specimens have been annotated by 
Larry Brown including most of Bill Carr’s 21 collections at TEX/LL. All 6 of his 
single taxa records have been examined and annotated. 

In most cases, nomenclature follows Jones et al. (1997) and Kartesz and 
Meacham (1999). Some recent literature sources were followed, especially some 


BROWN ET AL., FLORA OF THE BIG SANDY 


BIG THICKET NATIONAL PRESERVE 


ALABAMA - COUSHATTA .. wooovi? ~ cai 
INDIAN RESERVATION 


UPPER NECHES RIVER 
{ BEECH CORRIDOR UNIT 


CREEK UN 


HILLISTER(D 


WARREN (p 


Big Sandy | TURKEY 
Creek Unité! 


MENARD CREEK 
(CORRIDOR UNIT 


NECHES BOTTOM 
& JACK GORE 
BAYGALL UNIT 


\ 
‘N 


KOUNTZE aprove = 


ane! 


a 
i 


ry oO 


LOWER NECHES RIVER 
ORRIDOR UNIT 


2 
oO 
F 
< 
co 
z 
== 
a 


ROSIER UNIT “ I 


\Y 
LITTLE pee ISLAND 
RIOOR UNIT 


ORANGE CO 
‘| 


BEAUMONT UNIT 


Tain LAKE 


D: 


BEAUMONT 
— —_ 


\ 
JEFFERSDN CC 
| 
¢ 
cs 


Fic. 1. Location of Big Sandy Creek Unit, Big Thicket National Preserve, Texas. 


nomenclature in the Flora of North America volumes. In cases of multiple col- 
lections, no more than four are listed for each taxon. 


RESULTS 


Table 1 is an annotated list of the Big Sandy Creek Unit taxa. The following ab- 
breviations are used in Table 1 


CL=C. Liu. His specimen is at Rice University. 
CP = Charles Peterson. His two 2 aden are at SBSC. 
DR= David Rose 1? snecimens ar 


Be 


Hers 
GW = Geraldine yeeon Her specimens are at Rice University. 


708 BRIT.ORG/SIDA 22 


= 


1) 


LB = Larry Brown. His specimens are currently at Rice University except for 
those few that are indicated to be at SBSC. 

MM = Barbara and Michael MacRoberts. Their specimens are currently at Rice 

niversity except for one at SBSC. 

PR = Paul Rolling. His collections are a clear picture of Utricularia inflata from 
the Beaverslide Trail. There is no doubt as to its identification. One speci- 
men is at SBSC and a duplicate is currently at Rice University 

SEWP = Sandra Elsik and Warren Pruess. Their specimens are currently at Rice 


University. 

SJ = Stanley Jones. His specimens are oo at Rice University. 
WEE Bill Carr. His 21 specimens are at TEX/LI 

= exotic 

=see notes at end of list 
Those specimens listed as currently at Rice University are mounted and will 
be deposited at a yet to be selected major Texas herbarium. Except for the Stan- 
ley Jones specimens, they were collected under National Park Service Coopera- 
tive Agreement No. CA 14001004. 


ACANTHACEAE ANNONACEAE 

Hygrophila lacustris (Schlecht. & Cham.) Nees, SJ Asimina parviflora (Michx.) Dunal, SJ 16093; LB 
90;MM6929; LB 28616; GW 2582 28146, 28188 

Justicia ovata (Walter) Lindau var. lanceolata APIACEAE 


(Chapm,) R.W. Long, 5J 16240; LB 28170 Centella erecta (Lf) Fernald, S) 17408; LB 28629: 
Ruellia humilis Nutt., SJ 16340; LB 28760, 28219; MM 6256 


ae 
= 


Chaerophyllum tainturieri Hook. var dasycarpum 
ACERACEAE S.Watson, MM 6324 
Acer barbatum Michx., SJ 16242; LB 27804; GW — Chaerophyllum tainturieri Hook.var. tainturieri, LB 
2446 27843 


Acer rubrum L., SJ 16099; MM 6091; LB 27804, — Cicuta maculata L., GW 3502 
28147 *Cyclospermum leptophyllum (Pers.) Sprague ex 
AGAVACEAE Britton & Wilson, L 


Manfreda virginica (L,) Salisb. ex Rose, LB 28995, “/70sciadium beach DC, 5) 16429;GW : 04] 
Eryngium prostratum Nutt.ex DC. LB 2822 
28775 : 


Yucca louisianensis Trel., LB 27791;5J 16429 Hydrocotyle verticitata Thunb., SJ 16236 LB 
28657, 28243; GW 2606 
ALISMATACEAE Ptilimnium capillaceum (Michx.) Raf., SJ 16362; 
paecadie coat ifolius (L ey LB 28622 GW 2596 
Sagittaria la Willd., GW 2802 Sanicula canadensis L., MM 6275,6571;LB 28144 
AMARANTHACEAE GW 248. 
Froelichia floridana (Nutt.) Moq., MM 6709 Sanicula smallii Bickn., SJ 1618 
Spermolepis divaricata ie Raf. ex Ser., SJ 
ANACARDIACEAE | 16381: MM 6616 
us aromatica Aiton var. aromatica, LB 28669; lepis echinata (DC. Heller. MM 61444 


730 
Rhus copallinum L., SJ 16392; MM 6081 
sive up ae L., LB 28683 
tron radicans (L.) Kuntze, LB 27802 
Toxicodendron vernix (L.) Kuntze SJ 16188 


Thaspium trifoliatum (L.) A.Gray var.aureum Brit- 
ton, GW 2470 


Trepocarpus aethusae Nutt. ex DC., SJ 16254; LB 


BROWN ET AL., FLORA OF 


APOCYNACEAE 
Trachelospermum difforme (Walter) A. Gray, LB 
28790 


AQUIFOLIACEAE 
llex ambigua (Michx.) Torr., SJ 16233; LB 30452 
(SBSC) 


llex coriacea (Pursh) Chapm., LB 28955, 28148, 
28195; MM 6257 
llex decidua Walter, GW 2460, oe 2694, 2881 
llex longipes Chapm. ex Trel., SJ 16173; GW 2466, 
2882, LB 30460 (SBSC) 
flex opaca Soland., LB 28595, 28189, 28734; MM 


6100 
llex vomitoria Aiton, SJ 16095;:MM 6121 


ARACEAE 
Arisaema dracontium (L.) Schott, MM 6346; LB 
28346 


ee um (L oe LB 28242, 28621; 
MM 6258, GW 253 
16294; GW 


Peltandra Wane. (L ; Schott, SJ 1 
2464a, 2462a, 2641 


ARALIACEAE 
Aralia spinosa L.,SJ 16097; LB 28786; MM 6341 
*Hedera helix L., LB 28143 


ARECACEAE 
Sabal minor Jacq.) Pers., SJ 16079; MM 6073;LB 


ARISTOLOCHIACEAE 

Aristolochia reticulata Jacq., MM 6315;LB 28240 

Aristolochia serpentaria L., LB 28972; MM 6276, 
6549; GW 2017 


Aristolochia COMME LOISG Sims, GW 251 ] 


ASCLEPIADACEAE 

Asclepias ee Michx., SJ 16369; LB 
28374; MM 6699 

Asclepias perennis Walter, LB 30535 (SBSC) 

Asclepias tuberosa Woodson subsp. interior 


mee! cea variegata L., MM 6280; LB 28364 

pias verticillata L,LB 28779; GW 2016 
he. viridis Walter, MM 6310 
Matelea Ce ides (Engelm.) Woodson, MM 


656 
Moteea gaa! ipiens et are SJ 16182;LB 


MM 


Maree Roneenos (Walter) cone SJ 16415 
LB 28332,GW 2019 


PRESERVE 709 


ASPLENIACEAE 
aos ium platyneuron (L.) Britton, a & 
Poggenb., LB 28619, 28382; MM 6 


ASTERACEAE 
isiifolia L., LB 28672, 28719 
Ambrosia psilostachya DC., MM 6886; LB 28996 
Baccharis halimifolia L., LB 28950; GW 2735 
Berlandiera pumila (Michx.) Nutt. var. pumila, MM 
6285, LB 27801 


Ber! eo pumila (Michx.) Nutt. var. scabrella 
esom & B.L. Turner, LB 28431a; MM 6605 
oe bipinnata L., MM 6933:LB 28723 
Pie discoidea (Torr.& A.Gray) Britton, LB i 5 
ens frondosa L., MM 6934 8,27: 
ce tonia diffusa Elliott, MM 6907, 6727 
Chrysopsis mariana (L.) Elliott, LB 28943 
Chrysopsis pilosa Nutt., LB 28674, 28778; MM 
6614, 6611 
Cirsium horridulum Michx., MM 6152 
Conoclinium coelestinum (L.) DC., MM 6891: LB 
28937 


Conyza canadensis (L.) Cronquist var. canadensis, 
LB 28761 

Conyza canadensis (L.) Cronquist var. pusilla 
(Nutt.) Cronquist, LB 28415; MM 6708; GW 
2770 


Coreopsis lanceolata L., MM se 
Goleop sat Hnetona Nutt., MM 6 
Cr (Nutt) sr 16 29001; MM 


6704; WRC 10827 
+Croptilon hookerianum (Torr. & A. Gray) House, 


GW 2688 
+Diaperia candida (Torr.& A.Gray) Bentham, MM 
7049 


Echinacea pallida (Nutt.) Nutt. var. sanguinea 
(Nutt.) pe i R.D.Thomas, SJ 16402; LB 
28223a; MM 6 

Elephantopus oy, nianus Raeusch., MM 6946: 
WRC 10831 

Elephantopus nudatus A. Gray, MM 6896 

Flephantopus tomentosus L., LB 28598a, 28353; 


Erechtites hieraciifolia (L.) Raf.ex — LB 28960 
Erigeron annuus (L.) Pers., GW 262 
ee strigosus ee ex Willd., - 28153, MM 
084, 6107; GW 2 
at apillifoli om all) Lam., LB 28935 
Epaeaim lancifolium (Torr.& A.Gray) Small, LB 
28756 


= 


Fupatorium leucolepis (DC.) Torr. & A. Gray, MM 
6730, 6594 

Eupatorium perfoliatum L., LB 28613; MM 6344 

Fupatorium rotundifolium L.,LB 28427;MM 6729 

Eupatorium serotinum Michx., LB 28742, 28228 

ae ree (Torr. & A. ae Greene 
ex er & Britton, LB 28962, 

Facelis retusa (Lam.) Schultz-Bip., . er 


2490 
Gaillardia aestivalis (Walter) H.Rock var.aestivalis, 


LB 28680 

Gaillardia pulchella Foug. var. pulchella, MM 6236 

Gamochaeta coarctata (Willd.) Kerguelen, LB 
3045 1a (SBSC) 

Gamochaeta pensylvanica (Willd.) Cabrera, GW 
2585 


a 

Gamochaeta purpurea (L : - oe MM 6102;LB 
27786, 30537; WRC 1 

Helenium amarum ie - a var. amarum, 
MM 6545 


Helianthus angustif L.,LB 28947 
Helianthus hirsutus Raf.,LB 28402;MM 6750,6524 
Helianthus mollis Lam., LB 28676, 20400; MM 


Helianthus simulans E.E.Wats., LB 30877 
ber ee eet (Lam.) Britton & Rusby, 
LB 28932, 2 ae 0828 


— 


L., LB 28594; MM 6565 

Mi cnonenas artemisiifolius DC. var. 
artemisiifolius, SJ 16260;LB 28122; MM 6085 

*Hypochaeris microcephala (Sch. oe var, 
albiflora (Kuntze) Cabrera, MM 6 

lva annua L., GW 2052 

Krigia cespitosa (Raf.) Chambers, GW 2493 

Krigia virginica (L.) Willd, MM 6111;LB 27782 

Lactuca canadensis L.,LB 28409, 28378; MM 6518, 


6559 
Liatris acidota Engelm.& A. Gray, MM 6917 
Liatris Se he (Walter) Michx. var. elegans, MM 
747, LB 28984, 28762 
Liatris squarrosa (L.) Michx., var. hirsuta (Rydb.) 
r, LB 28990, 28684 
bon ise (L.) — var. squarrosa, MM 
6746; GW 2092, 
Wien ia ee. (L.) ci . 27807 
Oligoneuron nitidum yale : a en Small, LB 
28985, 28396; MM 669 
Packera glabella (Poir.) C. ee SJ 16237; MM 
6342; GW 2526 


BRIT.ORG/SIDA 22(1) 


Packera obovata (Muhl. ex Willd.) W. A.Weber & 
A.Love, LB 27805 

Pityopsis graminifolia (Michx.) Nutt., MM 6898, 
6080; LB 28772 

Pluchea camphorata (L.) DC.,LB 28934; GW 2785 

Pluchea foetida (L.) DC., LB 28710 

Pseudognaphalium obtusifolium (L.) Hillard & 
Burtt, LB 28934, 28673; MM 6716 

Pyrrhopappus carolinianus (Walter) DC., MM 
6313; SJ 15348 


Ratibida columnifera (Nutt.) Woot. & Standl., MM 
6543 


Rudbeckia hirta L., MM 6278, LB 28200; $/16353 
Smallanthus uvedalius (L.) Mack. ex Small, LB 
28/29a 


Solidago canadensis L., LB 28968; GW 2885 
Solidago ludoviciana (A. Gray) Small, LB 28749, 
28334, 27801a; MM 6294 
Solidago odora Aiton var. eee - 28942, 28660 
patie petiolaris Aiton., LB 
olidago rugosa Mill. nee asper (Aiton) 
Cronquist, LB 28946,22873 
Solidago rugosa Mill. subsp. rugosa, LB 28752,GW 
2424, 2423A;CL 901179 
*Soliva sessilis Ruiz & Pavon., GW 2476 
*Sonchus asper (L.) Hill, LB 29012 
*Sonchus oleraceus L., LB 2811 
Symphyotrichum dumosum (L.) Nesom, LB 


Roos, 


6a 
Symphyotrichum patens (Aiton) Nesom, LB 
28981, 27776 


Symphyotrichum pratense (Raf. 
6909 


<-> 


Nesom, MM 


Verbesina virginica L., LB 28780 

Vernonia missurica Raf., LB 30766 (SBSC) 

Vernonia texana (A. Gray) Small, MM 6725,6527; 
28731; GW 2731 

*Youngia japonica (L.) DC., LB 27839 

BERBERIDACEAE 

Podophyllum peltatum L., MM 6151 


BETULACEAE 

Alnus serrulata (Aiton) Willd., SJ 16144;LB 28172 

Betula nigra L., SJ 16297; MM 6328; LB 28249a 

Carpinus caroliniana Walter, MM 6538, 6148 

Ostrya virginiana (Mill.) K. Koch, SJ 16072, 16440; 
LB 28131;MM 6525 


ae 


ag ecnuanaine 
aL. LB 27834a 


BROWN ET AL., FLORA OF TH 


PRESERVE 711 


EC DIN SANUT 


meat Can ai ex Bureau, SJ 16422 
rd t) Warder ex Engelm., SJ 


16321; LB 27775 
BLECHNACEAE 
Wanaivard lata (L.) T. Moore, MM 6338; LB 


27846 
BORAGINACEAE 


y! oglossum virginianum fin, G 
Lithosperma tuberosum Rugel ex a 2213 
Myosotis macrosperma Engelm., GW 2491 


BROMELIACEAE 


(L.) L, MM 6936, 6330, 6755 
BUDDLEJACEAE 

Polypremum — L.,. MM 6566 
BURMANNIACEA 

AO SUED Nut LB 28782;CP 1041:DR 2373 
ta J.F.Gmel.) Mart., MM 6863 


CABOMBA 

Brasenia ae J.F.Gmel.,SJ 16291 
CACTACEAE 

Opuntia humifusa (Raf) Raf, SJ 16383; MM 6603 
CALLITRICHACEAE 

Callitriche heterophylla Pursh, SJ 16147;GW 2527 
CAMPANULACEAE 

Lobelia appendiculata A. - 6303;GW 2562 
Lobelia cardinalis L.,LB 2 

sere puberula Michx., ree 6878; GW 2822;LB 


28958, 28606 
Triodanis perfoliata (L.) Nieuwl., GW 2475 
*Wahlenbergia marginata (Thunb.) A. DC., SJ 
bs tee Sa NAKA | Z 


CAPRIFOLIACEAE 

*/ onicera japonica Thunb., MM 6263;LB 28156 

Lonicera sempervirens L., LB 27815; GW 2469 

Sambucus nigra L. subsp. canadensis (L.) R. Bolli, 
LB 28641 

Viburnum acerifolium L., SJ 16322 

Viburnum dentatum L., SJ 16396: MM 6074; 
LB28593; GW2465 

Viburnum nudum L. var. nudum, SJ 16286; LB 
28198 


Viburnum rufidulum Raf., MM 6093: GW 2040, 
3 


CARYOPHYLLACEAE 
*+Cerastium pumilum Curtis, MM 6143 (SBSC) 
072 


Gee 
Loeflingia squarrosa Nutt., MM 7 


+Minuartia muscorum (Fassett) Rabeler, SJ 16105 
+Silene subciliata B.L. Robins., LB 28599, 28626 


CELASTRACEAE 
Euonymus americanus L., MM 6071;LB 28163 


CHENOPODIACEAE 
Dysphania ambrosioides (L.) Mosyakin & Clem- 
ents, LB 28646 


CISTACEAE 

Helianthemum carolinianum (Walter) Michx, LB 
28235a, 27797; MM 6110 

Lechea mucronata Raf., LB 27783 

Lechea tenuifolia Michx.,SJ 16360;LB 28229: MM 
6609 


COMMELINACEAE 

ey ina say L.. MM 6601,LB 28385, 28169; 
SJ] 

one ei ee Le LB 29035; MM 6940 

Tradescantia reverchonii Bush, MM 6267 


CONVOLVULACEAE 
ae os is Michx., MM 6125; LB 


oe ae ie Slee LB 28333 

ipomoea lacunosa L., L 735 

— pandurata (L.) G. FW. Mey., LB 28429; 
MM 2753, 6617,6570 

Stylisma pickeringli A. Gray var. pattersonii 
(Fernald & B.G. Schub.) Myint, MM 6611 


CORNACEAE 

Cornus florida L., LB 28154; MM 6105; SJ 16394 

Nyssa aquatica L., SJ 16205, 16256; LB 28134, 
30455 (SBSC 


Nyssa biflora Walter, SJ 16317;LB 28322 

Nyssa sylvatica Marshall., MM 6429; LB 28759, 
30540 (SBSC); SJ 16087 

CUPRESSACEAE 

Juniperus virginiana L,MM 6114;LB 28598, 28356; 
SJ 16410 

Taxodium distichum (L.) L. C. Rich., SJ 16209, 
16339 


CUSCUTACEAE 

Cuscuta compacta Juss. ex Choisy, LB 30765 
(SBSC) 

CYPERACEAE 

Bulbostylis ee (Elliott) Fernald var 
ciliatifolia 
Bulbostylis Vee (Elliott) Fernald var 
coarctata (Elliott) Kral , SJ 16425 


712 


Carex abscondita Mack., SJ 16119; LB 30456 
(SBSC) 

Carex albicans Willd. ex Spreng. subsp. australis 
eal) JH. Rettig, LB 27835 


Carex chwein. 0566 
Carex atlantica Bailey subsp: capillacea (Bailey) 
Reznicek, MM 6244: LB 7 i 
Carex basiantha Steud., SJ 16 
Carex b/ ae ‘Dewey, SJ 161 . 
Carex Schwein, SJ 16115;LB 27820 
Carex ee mep nota Muhl. ex ae SJ 16121 


Car Schwein., SJ 16123 

Carex complanata Torr. & Hook., a 6279, 6529; 
C 17335 

Carex corrugata Fernald, SJ 16124; LB 27840 

Carex crebriflora Wieg., SJ 16041;LB 27836 

Carex debilis Michx.var. debilis, SJ 16117;LB 28181 

Carex digitalis Willd. var. asymetrica Fernald, SJ 


Carex festucacea schkuhr ex ane SJ 16044 

Carex floridana Schw 

Carex frankii Kunth, a ay 

Carex glaucescens Elliott, LB a 28266 

Carex intumescens Rudge, LB 27150 

Carex joorii Bailey, MM 6697; LB 28959a 

Carex leavenworthii Dewey, SJ 16103; 

Carex Seen Wahlenb. subsp. h i(F 

Stone, SJ 161 es = 2826/, 28201 

Carex one Mack., SJ 

Carex louisianica le 17352;S) 16212;LB 
28610, 2815 

Carex lupulina “en ex Willd.,SJ 17252;LB 28272, 
28209, 30563 (SB oe 

Carex lurida Wahlenb., LB 

Carex muehlenbergii ee ex Willd. var. enervis 
Boott, SJ 16043 

Carex oxylepis Torr.& Hook., SJ 16153; MM 6337 

Carex planispicata R. F.C. Naczi.SJ 16217, 16122 

Carex reniformis Small, SJ 16154 5262 

Carex retroflexa Muhl. ex Willd. SJ 16284 

iatula Michx., LB 30538 (SBSC) 

Face texensis Bailey, SJ 16104; MM 6130 

Carex triangularis Boeck., SJ 16118 

Carex tribuloides Wahlenb. var. a aa 
Clokey, SJ 16228, 16270;LB 2824 

Cyperus croceus Vahl, MM 6555;LB — 28387 

Cyperus echinatus Britton , SJ 16400 

Cyperus grayioides Mohlenbr. WRC 17705b 

Cyperus haspan L., LB 29015, 28729 


IAW 
ITlalQ) 


BRIT.ORG/SIDA 22(1) 


Cyperus hystricinus Fernald, MM 6756; DR 2364 
Cyperus plukenetii Fernald, LB 28671; MM 6720, 
6556; DR 2363 


Cyperus pseudovegetus Steud., SJ 16365; MM 
6698, 660 

Cyperus retroflexus Buckley, LB 28666; MM 6554; 
DR 2371 


Cyperus retrorsus Chapm., LB 28486, 28358 
Cyperus strigosus L., LB 28386, 28358 
Cyperus virens Michx., SJ 16337;LB 28155 
Fleocharis montevidensis Kunth, SJ 16349 
Eleocharis obtusa (Willd.) aoe 16424 
Eleocharis tortilis Schult., S 
Eleocharis tuberculosa a ens & JA. 
ultes, MM 6245,6578;LB 28323; =: 16363 
sie squarrosa Michx., MM 6596, 662 
ee odorata ve LB 28604; MM ae 
ica Elliott, SJ 16357 
sel eae corniculata (Lam.) A. Gray, MM 
6939; LB 28617 
Rhynchospora globularis (Chapm.) Small, var. 
globularis, SJ 16382;LB 28677;MM 6530,6599 
Rhynchospora glomerata (L.) Vahl, MM 6735; LB 
28430, 28418,28624 
Rhynchospora gracilenta A. Gray, MM 6887; LB 
28413; DR 2375 


Rhynchospora harveyi W. Boott, LB 28220; SJ 
16397 

Rhynchospora inexpansa (Michx.) Vahl, LB 28432, 
28324 


Rhynchospora mixta Britton, LB 28961, 28270, 
28340, 28248 
Rhynchospora pusilla Chapm. ex M.A. Curt. MM 


6574, 6576 

Rhynchospora rariflora Elliott, MM 6627;LB 28412 
5J 16 

Rhynchospora scirpoides (Torr.) A. Gray, DR 2376 

Scirpus cyperinus L. Kunth, LB 28745, 27848; MM 
6540 


Scleria ciliata Michx. var. ciliata, SJ 16201, 
MM 6292: LB 28237 
Scleria oligantha Michx., SJ 16280, 16126; MM 


16126; 


6325 
Scleria reticularis Michx., MM 6885 
Scleria triglomerata Michx., SJ 16423, 16224; LB 
28 28177 
CYRILLACEA 
Cyrilla oe L., LB 28179, 28241;MM 6591; 
SJ 16368 


BROWN ET AI 


PRESERVE 713 


DENNSTAEDTIACEAE 
Pteridium aquilinum (L.) Kuhn, MM 6139; LB 


DIOSCOREACEAE 
Dioscorea villosa L., LB 27833; SJ 16389 


DROSERACEAE 
Drosera brevifolia Pursh, MM 6094 
Drosera capillaris Poir., MM 6883,6575;LB 28416 


DRYOPTERIDACEAE 
Athyrium filix-femina (L.) Roth var. eae 
(Michx.) Farw. SJ 16275; MM 624 
Dryopteris ludoviciana (Kuntze) at GW 2600 
Onoclea sensibilis L., MM 6345; LB 28246, 27842 
arena Bees rode (Michx.) Schott, SJ 
16232; LB 28274, 27808 


EBENACEAE 
Diospyros virginiana LMM 6281;LB 28966, 28159 


ERICACEAE 
Rhododendron canescens (Michx.) Sweet, SJ 


16329 

Rhododendron viscosum (L.) Torr., SJ 16289; LB 
2895, 28279,27850 

Vaccinium arboreum Marshall, SJ 16214; LB 

28664, 28145; MM 6291 

Vaccinium corymbosum L., LB 28428; 28129; MM 
6137; GW 201 

Vaccinium stamineum L., SJ 16066; MM 6320 


ERIOCAULACEAE 

Er iocaulon te Croke Koern., MM 6592: SJ 16372 

EUPHORBIACEAE 

Acalypha oe A. Gray, MM 6872, 6757; LB 
28953; GW 2768 


Acalyph cane idea Raf., GW 2603 

Chamaesyce cordifolia (Elliott) Small, LB 28675, 
28768; MM 6618 

Cnidoscolus texanus (Muell.-Arg.) Small, MM 
6269; SJ 16 

Croton argyranthemus Michx., MM 6309; LB 
28224; GW 2576 


Croton Michx., MM 6718 

Croton g Danahiostis L. var. septentrionalis Muell.- 
Arg. “MM 6695, GW 2602; LB 28722, 28370 

Croton michauxii G. L. Webster, GW 2687; MM 


6619 
Euphorbia corollata L., LB 27770; MM 6544; GW 
2045 


pede ou jensis Walter, LB 28607; GW 


pre ra urinaria L.,LB 27927; MM 6945 

Sebastiania fruticosa (W. pee Fernald, SJ 
16152; MM 6327; LB 28187, 27830 

Stillingia sylvatica L., SJ 16367; a 6711 

Tragia smallii Shinners, GW 2577 

Tragia urticifolia Michx., MM 6306,6714,6548;LB 


28379 
*Triadica sebifera (L.) Small, SJ 16446 


FABACEAE 
*Albizia julibrissin Durazz., SJ) 16445; LB 29006, 
27771; MM 6524 


Baptisia bracteata Muhl. ex Elliott var laevicaulis 
(A. Gray ex Canby) D. Isley, SJ 16106 

Baptisia bracteata Muhl.ex Elliott var.leucophaea 
(Nutt.) Kartesz & Gandhi, MM 6132 

pe nuttalliana Small, MM 6316, 6133 

entrosema virginianum (L.) Benth., LB 28381 

a is canadensis L.var. canadensis, MM 6150;LB 
28681; SJ 28381 

Chamaecrista fasciculata (Michx.) Greene, MM 

751; LB 28681 
Chamaecrista nictitans (L.) Moench, LB 28711 


28658 

Clitoria mariana L.,LB 28590, 28757, 28339: MM 
6553 

Crotalaria sagittalis L.,SJ 16220; MM 6693, 6551; 

B 28327 

Dalea phleoides (Torr. & A. Gray) Shinners, MM 
6748 

Desmodium ciliare a ex Willd.) DC.,LB 29032, 
28376; GW 2714 


Desmodium ie Hi ens DC., LB 28734, 
GW 2773 


ae\ ae m yk as LB 29030 
DC,,LB 2 


bt, 


OUlUSUT Ti 


Demoaon paniculatum DC,, a 0;GW 2422 
Desmodium rotundifolium DC., LB 28668; GW 


2803 
Erythrina herbacea L., SJ 16322; MM 6288; LB 


28223 

Galactia volubilis (L.) Britton, LB 28769; MM 6526, 
6552; GW 2578 

Gleditsia aquatica Marshall, LB pe 

Indigofera suffruticosa Mill. GW 

*Kummerowia striata Mill., LB oars 28925 

*Lathyrus hirsutus L.,GW 2011 

Lespedeza hirta Elliott, MM 6925;LB 29000 


Lespedeza procumbens Michx., LB 28397, 28380, 
440 
Lespedeza repens (L.) Barton, LB 28758, 28231a; 


Lespedeza stuevei Nutt., LB 28767, 29019; GW 
2/94 


Lespedeza virginica (L.) Britton, LB 28751, 28232; 
G 6 

Mimosa hystricina (Small ex Britton & Rose) B.L. 
Turner, SJ 16258;LB 28216 

Rhynchosia difformis (Elliott) DC. LB 28750, 
28667,2 GW 2413 

Rhynchosia latifolia Nutt. ex. Torr. & A. Gray, GW 


2565, 2/33 
Rhynchosia reniformis DC.,LB 28406, 28235; MM 
6270,6314 
pil mari joloes (L.) a LB 28642; GW 2049 
(L.) Irwin & Barneby, GW 2715 
(L.) eae GW nee 
+Sophora affinis Torr. & A. Gray, G 
Strophostyles umbellata (Muhl. ex vie Britton, 
8781, 28754 
Stylosanthes biflora (L.) Britton, Sterns, Poggenb., 


FC it tol 


Tephrosia onobrychoides Nutt., MM 6283; LB 
eas GW 2463 

Tep! iniana (L.) Pers.,SJ 16391;LB 28661, 
2821; 7 MM 6301 

He ea ium eee Schreb., MM 6146;SJ 16354 

Sibthrop, LB 27845b 

*Trifoli um incarnatum L., SJ 16062; MM 6323 

*Trifolium repens L., SJ 1 —_ 

*Vicia sativa L. MM 614 

Wisteria frutescens (L.) ae SJ 16207; LB 28174, 
28171, 28175 

FAGACEAE 

Castanea pumila (L.) Mill. SJ 16314 

Fagus grandifolia Ehrh., MM 6101; SJ 16279; LB 


= 


27781 

Quercus alba L., LB 28125; MM 6602, 6127; SJ 
16075 

Quercus falcata Michx., MM 6348, 6523; LB 28600; 
SJ 16304 

Quercus hemisphaerica W. Bartram ex Willd., MM 


6532; SJ 16110; LB 28777, 28713 
] artram, MM 6274,6568,6567; 


Quercus laurifolia Michx., LB 27844, 27827a, 
30560 (SBSC) 


BRIT.ORG/SIDA 22(1) 


Quercus lyrata Walter, LB 28748 
ercus margaretta Ashe ex Small, MM 6318 
6311 

Quercus marilandica Muench., MM 6090; SJ 

LB 27778, 28665 
Quercus michauxii Nutt., LB 28649: MM 6078, 
+ SJ 16084 

Quercus nigra L., MM 6077, 6261; SJ 16098; LB 
28183 

Quercus pagoda Raf., SJ 16169 

Quercus phellos L.,LB 28117,27733;MM 6268; S5 


16215 
Quercus stellata Wangenh., SJ 16108;LB 29005 
GENTIANACEAE 
ne ice, s(L ae CP 
t.MM ae . 28239 


GQUaUG CANIp 

GERANIACEAE 

Geranium carolinianum L., SJ 16068 

GROSSULARIACEAE 

Itea virginica L., SJ 16068; LB 29013, 28133; MM 
6243 

HALORAGACEAE 

*Myriophyllum aquaticum (Vell.) Verdc. SJ 16292; 
GW 3345 


Id Cri 


HIG OOS, 


Proserpinaca palustris L.var.crebra F 
LB 28741 


Proserpinaca palustris L.var.amblyogona Fernald, 
LB 28158 


HAMAMELIDACEAE 

Hamamelis virginiana L., SJ 16307; LB 28176 

Liquidambar styraciflua L., SJ 16073, 16441; MM 
6120 


HIPPOCASTANACEAE 
Aesculus pavia L., MM 5234 


HYPERICACEAE 
st drummondii (Grev, & Hook.) Torr. & A. 
B 28922, 28765; MM 6613; GW 2793 

eee galioides Lam., LB 28422; MM 6587 

Hypericum gentianoides (L.) Britton, Sterns, 
Poggenb., MM 6892, 6621; . 28988, 28764 

z,MM 6098 

ae uit LLB 28417, ae 60; MM 6625 

Triadenum walteri (J.G.Gmel.) Gleason, MM 6931, 
6953; LB 29016; GW 2821 

IRIDACEAE 

Alophia drummondii (Graham) R. C. Foster, MM 
6516; LB 28392, 28234; SJ 16416 


H\ peri wim Ay peri oides (L.) Cra 


PRESERVE 715 


BROWN ET AL., PLUAA VE TTT 


Herbertia lahue (Molina) Goldblatt, MM 6253; SJ 
16190 
Iris ssid icaulis Raf., SJ 16251 


hium anaustifolium Mi IL, G 


Sahm etiam Bae eat ae 6263; 
GW 2495 


Sisyrinchium sagittiferum E. P. Bicknell, MM 614 
WRC 17353 


JUGLANDACEAE 
Carya cordiformis (Wangenh.) K. Koch, SJ 16282 
Carya glabra (Mill.) Sweet, LB 28623, 28679,28373; 


7 
Carya texana Buckley, LB 28391; MM 6271 
Carya tomentosa (Lam. ex Poir.) Nutt., SJ 16287; 
LB 27772; GW 2447, 2448 
Juglans nigra L., SJ 16393; LB 28609; MM 6349 


JUNCACEAE 

Juncus acuminatus Michx., LB 30561 (SBSC) 
Juncus capitatus Weigel, SJ 1604 

Juncus coriaceus Mack., MM 6935,6528;LB 28632, 


28967 
Juncus dichotomus Elliott, SJ 16385; MM 6604 
Juncus diffusissimus Buckley, SJ 16296 
Juncus effusus L., SJ 16138; MM 6254; LB 28207, 


Juncus marginatus Rostk, SJ 16346 

Juncus repens Michx., MM 6944; LB 27851 

Juncus scirpoides Lam., LB 28421; MM 6607 
Juncus tenuis Willd., LB 28161; SJ 16405 

Juncus trigonocarpus Steud., LB 28161;DR 2366 

Juncus validus Coville, SJ 16352, 16406;LB 28328 


LAMIACEAE 
Hedeoma hispida Pursh, MM 6624 
Hyptis alata (Raf.) Shinners, MM ak 
cae rubellus Moench, MM 694 
COPUS VIFGINICUS ls GW oe a 2966 
Moe fistulosa L.GW 2 
Monarda punctata L. MM i 7,6705;LB 28618 


GW 2 

*Perilla frutescens (L.) Britton, LB 28643 

Prunella vulgaris L., MM 6305 

Pycnanthemum albescens Torr. & A. Gray, LB 
28736, 28755; MM 6308, 6707 

Pycnanthemum tenuifolium Schrad., SJ 16432; 
MM 6703 


Salvia azurea Michx. var. grandiflora Benth., MM 
6926; LB 29023 
Salvia lyrata L., MM 6117;LB 28645, 27809 


— 


Scutellaria ees Engelm. & A. Gray, LB 
2 


(e lla Ili i; Muhl.e 
“LB 28365, cee 

Scutellaria integrifolia L.,LB i 

Scutellaria parvula Michx., GW ast 17333 

hed fenuONG Willd., GW 272 

L.,LB — GW 2775 

iichosteme setaceum Houtt., MM 6924a; LB 

29022 


ve SJ 16276; 


LAURACEAE 

Persea borbonia (L.) Spreng., MM 6273, 6087; LB 
28126, 28390 

Persea palustris (Raf) Sarg., LB 27847, 28325; SJ 
16397 


Sassafras albidum (Nutt.) Nees, LB 28639; MM 


poate LARIACEAE 
inguicula pumila Michx., GW 
uu cornuta Michx., LB ee (SBSC) 
tricularia gibba L., LB 28717 
icularia inflata Walter, PR s.n. 
icularia juncea Vahl, MM 6585 
Utricularia subulata L., GW 2487 
LILIACEAE 
Alli d L.var. canadense, MM 6282;GW 


2537 

Allium ae L. var. mobilense (Regel) 
Ownbey, SJ 16197 

Hypoxis curtissil ee MM 6343; GW 2522; LB 
28175a; SJ 16235 

Nothoscordum bivalve (L.) Britton, MM 6095 

LINACEAE 

Linum medium (Planch.) Britton var. texanum 
(Planch.) Fernald, MM 6533;LB 28424, 28218 


2574 
Linum striatum Walter, GW 2598, 2719; MM 6519 


LOGANIACEAE 
Gelsemium sempervirens (L.) Aiton f, MM 6113 
SEWP 2005 


Mitreola petiolata (J.F. Gmel.) Torr. & A. Gray, LB 
29017, 28739; GW 2780 

Mitreola sessilifolia (J.F. Gmel.) G.Don, LB 28431; 
MM 6626; 2695 

LYCOPODIACEAE 

Lycopodiella appressa (Chapm.) Cranfill, MM 
6890, 6579 


Pseudolycopodiella caroliniana (L.) Holub var. 
caroliniana, MM 6582; GW 2712 


LYGODIACEAE 
*Lygodium japonicum (Thunb. ex Murray) Sw 
MM 6317;SJ 16211;WRC 10845 


LYTHRACEAE 
*Cuphea pele le 
6298, 674 3 
*Cuphea ss Cham.& Schlecht., MM 6266; 
LB 28393; GW 2512 
otala ramosior (L.) Koehne, GW 2782 


MAGNOLIACEAE 
Magnolia grandiflora L., LB 27818,28196; MM 
6149; SJ 16070 


(Jacq.) J.F. Macbr., MM 


Magnolia virginiana L., MM 6248; LB 27813; SJ 
1637] 


MALVACEAE 
Mal j t Dill.ex.Cav.var.drummondii 
(Torr.ex A. Gray) Schery, LB 28345; MM 6719 
Sida rhombifolia L., MM 6752;LB 28637 

Sida spinosa L., GW 2725 


MAYACACEAE 

Mayaca fluviatilis Aubl.,, GW 2507, 2788 
MELASTOMATACEAE 

Rhexia mariana L., MM 6608;LB 28330;GW 2597, 


2969 
Rhexia virginica L., LB 28728; MM 6733 
MELIACEAE 
*Melia azedarach L., LB 28964 
MENISPERMACEAE 
Cocculus carolinus (L.) DC., LB 28203 
MOLLUGINACEAE 
Mollugo verticillata L., MM 6606; SJ 16377 
MONOTROPACEAE 
Monotropa uniflora L., LB 28372 


MORACEAE 
Morus rubra L., MM 6123;LB 28776;SJ16438;GW 
3335 


MYRICACEAE 

Morella caroliniensis (Mill.) Small, MM 6237, LB 
28956, 28784, 28182 

Morella cerifera (L.) Small, MM 6340; LB 28130, 
28247; GW 2707 


BRIT.ORG/SIDA 22(1) 


NYMPHAEACEAE 
bi lutea (L.) Sm. subsp. advena (Aiton) 
artesz & Gandhi, LB 26168 


OLEACEAE 
Chionanthus virgin 
SJ 16089 

Forestiera ligustrina (Michx.) Poir., GW 1944, 2421, 
2515; LB 27817 

Fraxinus americana L., MM 6319; GW 2708; LB 
28106; SJ 16365 

Fraxinus caroliniana Mill., 
(SBSC) 


icus L., MM 6118; LB 28137, 


SJ 16203; LB 30565 


LGN ll lait Marshall., MM 6076 
r., LB 28945; SJ 16251 


CNAGRACERE 
a alternifolia L.,LB 28331; GW 2799 
oc a pl Walter GW 2798; LB 28724; 


eae are Walter, GW 2720, 2967; LB 
28744 


Ludwigia hirtella Raf; LB 28328; MM 6585, 6734 

Ludwigia leptocarpa (Nutt.) Hara, GW 2968 

Ludwigia linearis Walter, MM 6897 

Ludwigia pilosa Walter, LB ae ie 

Oenothera laciniata Hill, GW 

Oenothera linifolia Nutt., MM pe 

Oenothera speciosa Nutt., MM 6260; SJ 16344 

OPHIOGLOSSACEAE 

Botrychium biternatum (Sav.) Underwood, MM 
6910; LB 27831 

Botrychium virginianum (L.) Sw., MM 6331,6147 
GW 1945 


ORCHIDACEAE 

Corallorhiza wisteriana oe aa 2007 

Listera australis Lindl., SEWP 

Platanthera clavellata ne ne LB 28268;GW 
2563 


Plata th 


ta (Michx.) Lindl., GW 2701 
Spiranthes cernua (L.) L.C. Rich., MM 6952; GW 


3449 
Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) 
Luer, MM 6620 
Spiranthes praecox (Walter) S.Watson, SJ 16259 
Spiranthes tuberosa Raf., LB 28382;MM 6573 
OROBANCHACEAE 
Epifagus virginiana (L.) W. Bart., LB 28271 


BROWN ET AL., FLAA VET 


OSMUNDACEAE 
Osmunda cinnamonea L., MM 6242;LB 28190 
Osmunda regalis L.var spectabilis (Willd.) A.Gray, 
MM 6259; LB 28123, 28206, 27855 


OXALIDACEAE 

Oxalis corniculata L. var. wrightii (A. Gray) B. L. 
Turner, MM 6096 

*Oxalis debilis Kunth var. corymbosa (DC.) 
Lourteig, $J 16231 

Oxalis lyonii Pursh, (T) MM 6135;WRC 17337 


PASSIFLORACEAE 
Passiflora lutea L., MM 6293, 6742; LB 28939, 
28338 


— 


PHYTOLACCACEAE 
Phytol icana L., LB 28140 


PINACEAE 

a ead Mill. MM 6893, 6082; LB 28949; SJ 
q I, MM 6070; SJ 1631 

ate taeda L., MM 6083; SJ 16257 


PLANTAGINACEAE 
Plantago aristata Michx.,SJ 16295;LB 28236; MM 
6623 


PLATANACEAE 
Platanus occidentalis L., MM 6246; SJ 16434; LB 
27816; GW 2046 


POACEAE 
pie is elliottiana J.A.Schultes, GW 2501 
tis hyemalis (Walter) Britton, Sterns, 
*Poggenb, 
A legans inth, MM 6235;LB 28212 
aneronocen gerardii Vitman, LB 29033, 28999 
Andropogon glomeratus (Walter) Britton, Sterns, 
Poggenb. LB 28965; GW 2784 
Andropogon gyrans Ashe var.gyrans, WRC 17339 
Andropogon ternarius Michx., MM 6874,6916;GW 
2807; LB 28982 


Andropogon virginicus L.var. virginicus, LB 29027, 
28997, 28941, 28936 

Aristida lanosa Muhl. ex Willd., MM 6927; LB 
29002; WRC 10829; DR 2367 

Aristida longispica Poir. var. geniculata (Raf.) 

rnald, MM 6904 

Aristida se Poir. var. longispica, LB 28926; 

GW 2789, 2 
ristida ah aes GW 2791 


PRESERVE 717 


Aristida purpurascens Poir. var. purpurascens, MM 
6871; LB 29004, 28991, 28931 

Arundinaria gigantea (Walter) Muhl., MM 6072; 
LB 28650, SJ 16120; SEWP 2001 

Axonopus fissifolius (Raddi) Kuhlm., LB 28359, 
28611; GW 2705 

*Briza minor L.,MM 6079; GW 2611 

*Bromus japonicus Thunb. ex Murr., SJ 16359 

Cenchrus spinifex Cav., LB 28363; MM 6615; GW 
2/04 


Chasmanthium lfatifolium (Michx.) Yates, SJ 
B 28350, 28151; MM 6743 
Chasmanthium laxum (L.) Yates var. laxum, LB 


30 
Chasmanthium laxum (L.) Yates var. sessiliflorum 
(Poir.) Wipff & S. Jones, MM 6722; LB 28714, 

28348 


Dichanthelium aciculare (Desv. ex Poir.) Gould & 
Clark SJ 16060, 16107;LB 28238 
Dichanthelium aciculare (Desv. ex Poir.) Gould & 
Clark subsp. angustifolium (Elliott) Freckman 
& Lelong, LB 28367 
Dichanthelium acuminatum (Sw.) Gould & Clark 
var.acuminatum, SJ 16427;LB 28335 
Dichanthelium boscii (Poir.) Gould & Clark, LB 
28273, 28663, 28351, 28149 
Dichanthelium commutatum (Schultes) Gould 
subsp , MM 6239; SJ 16064, 
i i a ent (SBSC) 
Dichanthelium commutatum (Schultes) Gould 
ubsp. equilateral (Scribn.) Freckman & 
Lelong, LB 28589 
Dichanthelium dichotomum (L.) Gould var. 
dichotomum, MM 6299; LB 28975, 28628; SJ 
16171 
Dichanthelium dichotomum (L.) Gould subsp. 
microcarpon (Elliott) Freckman & Lelong, LB 
28166, 28164 
Dichanthelium ensifolium (Baldw. ex Elliott) 
ould subsp. ensifolium, LB 29038; SJ 16308 
ohana laxiflorum (Lam.) Gould, LB 27788; 
WRC 1 


See 


Sein oligosanthes (Schult.) Gould, var. 
oligosanthes, SJ 16184 

Dichanthelium oligosanthes (Schult.) Gould var. 
scribnerianum (Nash) Gould, LB 28369 

Dichanthelium oligosanthes (Schult.) Gould, WRC 
17340 


Dichanthelium ovale (Elliott) Gould & C.A. Clark 


subsp. villosissimum (Nash) Freckman & 
Lelong, SJ 16263, 16195; MM 6277; WRC 
17341 


Dichanthelium polyanthes (Schult.) Mohlenbr,, LB 
28384, 30567 (SBSC), 30573 (SBSC); SJ 16404 

Dichanthelium ravenelii (Scribner & Merr.) Gould 
& Clark, MM 6290; SJ 16435 

Dichanthelium scoparium (Lam.) Gould, SJ 16407; 
LB 28405; MM 6612 

Dic. pe ium sphaerocarpon (Elliott) Gould, MM 
6 3; SJ 16265 LB 28630 

See ane wrightianum (Scribn.) Freckmann, 
LB 28433; MM 6583 


Digitaria ciliaris (Retz.) Koel., LB 28933, 28712 
Digitaria cognata (J.A. Schultes) Pilger var. 
LGW 2815 
Digitaria filiformis (L.) Koel. var. filiformis GW 2806 
Digitaria filiformis (L.) Koel. var villosa (Walter) 
Fernald, MM 6876; LB 29026; GW 2418 
*Digitaria violascens Link, LB 29009 


Elymus virginicus L., LB 28208, 28732; MM 6535; 
SJ 16444 


Eragrostis elliottii S.Watson, GW 2812 
Eragrostis hypnoides (Lam.) Britton, Sterns, 
Poggenb 19b 


— 


Eragrostis intermedia (Lam.) Britton, Sterns, 
g 894 


Eragrostis refracta (Muhl.) Scribn., LB 28989 
— sree (UJ. Presl) subsp. oxylepis 

(Tor Koch, MM 6696;LB 28774;GW 2797 
(Pursh) Steud., MM 6915, 
983 


oe spectabilis 
6869, 6902; LB 28 

Eustachys petraea (Sw.) Desv., MM 6903 

Gymnopogon ambiguus (Michx.) Britton, Sterns 

nb., MM 6865; GW 2808; DR 2368 

Leersia virginica Willd., LB 28971, 28627 

*Lolium perenne L., MM 6322;SJ 16350 

Melica mutica Walter, MM 6108;LB 28173, 27832 

Oplismenus hirtellus (L.) Beauv., LB 28648: MM 
6938 


Panicum anceps Michx., MM 5757, 5646; LB 


29008, 28974 

Panicum brachyanthum Steud., MM 6912; LB 
68980, 68929; WRC 10832 

Panicum gymnocarpon Elliott, LB 28973, 28740 

Panicum rigidulum Bosc ex Nees, LB 2874 

Panicum verrucosum Muhl., MM 6881,6950; LB 
29037, 2901 | 

*Paspalum notatum Fluegge, MM 6875, 6537;LB 
28603; GW 2585a 


BRIT.ORG/SIDA 22(1) 


Paspalum plicatulum Michx., MM 6864, 6731;LB 
28199; SJ 16356 

Paspalum praecox Walter, MM 3933, 3719, 3937 

Paspalum setaceum Michx., GW 2419: LB 
2403,28928; MM 6724 

*Paspalum urvillei Steud., MM 6895, 6541; LB 
28221, 29010 


Phalaris angusta Nees ex Trin., SJ 16255 
Piptochaetium avenaceum (L.) Parodi, MM 6329; 
SJ 16378 
*Poa annua L., MM 6142; GW 2497, 2498 
Poa autumnalis Muhl. ex Elliott, LB 27829 
Saccharum ‘sania (Walter) Pers., MM 6880 
Se ne scoparium (Michx.) Nash var. 
eee Gould, LB 28998; GW 2814 
setae paraiora (Poir.) Kerguelen, LB 28361 
sh, MM 6919 
Sphenophols filiformis (Chapm.) Scribn., WRC 
7336 


nes 


Sorghastn || ttii (Mo hr) Na 


Sph en resect atnyes) Rydb., SJ 16229 
Sphenopholis obtu hx.) Scribn., MM 6300 
Sporobolus niet inus ee A.S.Hitche.,, MM 
peed LB ian gis GW 2813 
j R.Br, LB 28930 
Sorat us junceus (Beauv) Kunth, LB 28993:MM 
0; DR 2369 


—_ 


Steinchisma hians (Elliott) Nash, SJ 16298 

Tridens flavus (L.) ead var. chapmanii (Small) 
Shinners, WRC 1082 

Tridens flavus (L.) Tee flavus, LB 28634;GW 
2426, 2819/2783 

Tridens strictus (Nutt.) Nash, LB 28904 

POLEMONIACEAE 

Phlox nivalis Lodd. ex Sweet subsp. texensis 
Lundell, MM s.n.;WRC 17332 


POLYGALACEAE 
olygala mariana Mill, LB 28215; SJ 16206 
Polygala polygama Walter, MM 6296;LB 28116 


POLYGONACEAE 
Brunnichia ovata (Walter) Shinners, MM 6951:LB 
28138 


Polygonum punctatum Elliott, MM 6930; LB 
28656; GW 2883, 2778a 
Polygonum eral L., LB 28656a 


POLYPODIACEA 
Pleopeltis por ypodioide ANTAIEWs & rhe oi 
sub sp ndrews & 


pARSER tds 


Windham, LB 28407, 37795 


BROWN ET AL., FLORA OF TH 


C DIN SANUT 


POTAMOGETONACEAE 

Potamogeton diversifolius Raf., SJ) 16273; LB 
28682; 

Potamogeton pulcher Tuckerman, GW 2561 


PRIMULACEAE 
An Ili is L., MM 6089 
Anagallis minima (L.) Krause, LB 277 
Samolus valerandi L. subsp. a iss (Raf.) 
Hultén, LB 29040; SJ 1624 


RANUNCULACEAE 
Clematis reticulata Walter, GW 2734; MM 6610 
Ranunculus hispidus Michx. var. nitidus (Chapm. 
T.Duncan, . 27841; GW 2734 
*Ranunculus parviflorus L., GW 2517 
Ranunculus pusillus Poir., GW 2480 
ecurvatus Poir., GW 3351 


RHAMNACEAE 

Berchemia scandens (Hill) K. Koch, MM 6103; LB 
278615) 16159 

Rhamnus caroliniana Walter, MM 6122:LB 28633, 
27799, 30572 (SBSC) 


ROSACEAE 
Agrimonia microcarpa Wallr., GW 2769,3425; MM 
6721 


ro 


Agrimonia rostellata Wallr., LB 28732; MM 6334 

Crataegus marshallii Egglest., MM 6726;LB 28970, 
27825a; SJ 16074 

Crataegus opaca Hook. & Arn., SJ 16078 

Crataegus spathulata Michx., SJ 16096;LB 28644, 
2/ ] 


Crataegus uniflora Muench., GW 2014 

Crataegus viridis L. GW 1947a 

Prunus caroliniana (Mill.) Aiton, LB 28951 

Prunus gracilis Engelman & A.Gray, MM 6284; LB 
27785 


Prunus serotina Ehrh., MM 6119;LB 28785, 28269; 
SJ 16067 

Prunus umbellata Elliott, LB 27800; SJ 16172 

Rubus argutus Link, SJ 16155;LB 28788; GW 2706, 


2457 
Rubus trivialis Michx., MM 6116 


RUBIACEAE 

Cephalanthus occidentalis L., MM 6932, 6593 
Diodia teres Walter, MM 6694; LB 2840 

Galium pilosum Aiton, GW 2573; MM 6326 
Galium tinctorium (L.) Scop., LB 27806; GW 2483 
Galium triflorum Michx., SJ 16283; LB 28789 


BIG THICKET NATIONAL PRESERVE 719 


Galium uniflorum Michx., SJ 16278; LB 30460a 

BSC) 

Houstonia micrantha (Shinners) Terrell, WRC 
17334 

Houstonia pusilla Schoepf, MM 611 

Mitchella repens L., MM 6247, 6189; LB 27803; SJ 
161 ee 

Oldenlar L., GW 2713,2790 

penroden pentane (K.Schurm.) Vatke, LB 28608; 

2374 


— 
W 


aL, LB 28924 
*Sherardia arvensis L., MM 6136 
Stenaria nigricans (Lam.) Terrell, LB 28368, 28222; 


RUTACEAE 
*Poncirus trifoliata (L.) Raf.,SJ 16246;LB 28652 
Zanth herculis L.,LB 28185 


| 'PaiVial 


ylun 
SALICACEAE 
Salix nigra Marshall. SJ 16142 


SAPOTACEAE 
Sideroxylum lanuginosum Michx.subsp.oblongi- 
folium (Nutt.) T.D. Pennington, SJ 16306 


SARRACENIACEAE 
Sarracenia alata Wood, MM 6589 


SAURURACEAE 
Saururus cernuus L., LB 28620, 28142a; MM 6943 


SAXIFRAGACEAE 
Lepuropetalon spathulatum Elliott, LB 27768 


SCROPHULARIACEAE 

Agalinis fasciculata (Elliott) Raf, LB 28948, 28923 
+Agalinis filifolia (Nutt.) Raf, MM 6921,6922,6901 
Agalinis purpurea (L.) Pennell, LB 29029; GW 2879 
Aureolaria flava (L.) Farw.var macrantha Pennell, 


639 
Aureolaria grandiflora (Benth.) Pennell, MM 6751; 
LB 28591; GW 2736 
Aureolaria pectinata (Nutt.) Pennell, GW 2880 
Bacopa caroliniana (Walter) Robins. LB 30562 
(SBSC) 


Gratiola neglecta Torr, GW 2526 
Gratiola pilosa Michx., GW 2696; LB 29036; MM 
6581 


Gratiola Virgil al 1d L, GW 2485, 2525), 2604 
Lindernia dubia (L.) Pennell, SJ 16146; LB 28720; 


GW 2605 
Mecardonia acuminata (Walter) Small,GW 2781; 
LB 29028, 28230 


720 


Mecardonia procumbens (Mill.) Small, GW 809 
Micranthemum umbrosum (J.G.Gmel.) Blake, SJ 
16269; 28592 
Pe nctemon iflary) 
16077; LB 28227 
Scoparia dulcis L., MM 6692; LB 28357, 28225 


SELAGINELLACEA 
Selaginella apoda (L ‘ Spring, MM 6255;GW 2508; 


Pennell, MM 6307, 6088; SJ 


SMILACACEAE 
Smilax one: nox L., MM 6691 
Smila ca Walter, LB 28977 


in Lunplia L., MM 6250, se : 28193:SJ 


Smilax pumila Walter, MM 6086; SJ 1616 
Smilax rotundifolia b., MM 6238: SJ 16436; LB 


27779 

Smilax smallii Morong, MM 6240; LB 28938, 
28233; SJ 16158 

Smilax tamnoides L.,SJ 16202 


SOUEN RSENS 

Physali phylla Nees, i 6286; GW 2801 
Physalis pubescens L., GW 2599 

al pau Nutt. MM 5369, 5404; GW 2660 
M 6723;LB 28337 
*Solanum capsicastrum Link ex Schauer, GW 


inense L 


2047 
Solanum ptycanthum Dunal, MM 6937;SJ 16248 


SPARGANIACEAE 

Sparganium americanum Nutt., SJ 16293: LB 
30564 (SBSC) 

STYRACACEAE 

Halesia diptera Ellis, SJ 16412;LB 27825 

Styrax grandifolius Aiton, LB 28128, 28124,28127 


SYMPLOCACEAE 
Symplocos tinctoria (L.) L.Her, SJ 16318;LB 28783, 
28277 


THELYPTERIDACEAE 

Thelypteris kunthii (Desv.) Morton, MM 6339; SJ 
16227; LB 26244, 26244 

TILIACEAE 

Tilia americana L. var. caroliniana (Mill) Castigl., 
LB 27828; GW 2012, 2048 


ULMACEAE 

Celtis laevigata Willd., GW 2513b,2514c 

Celtis tenuifolia Nutt,, MM 6522:LB 30534 (SBSO), 
30560 (SBSC) 


— 


at 


BRIT.ORG/SIDA 22(1) 


Planera aquatica J.F. Gmel., LB 28180; SJ 
16204,6414; GW 2042 

Ulmus alata Michx., MM 6706, 6092; GW 2415:S) 
16439 

Ulmus americana L., SJ 16132,5J 16140;LB 28211; 
GW 2425 


URTICACEAE 
Boehmeria cylindrica (L.) Sw., LB 28344, 28136; 
MM 6744 


VALERIANACEAE 
Valerianella radiata (L.) Dufr., MM 6333 


VERBENACEAE 
Callicarpa americana L., SJ 16442;LB 28347;MM 


a 


6 9 
Glandul sg canadensis (L.) Nutt. MM 6115; LB 


2840 

es orem ) Greene, SJ 16431 

Vell., SJ 6908 

Veibene halei Small MM 6308; LB 28236a; GW 
2689 


na brasili 


*Verbena rigida Spreng., MM 6542 
*Vitex agnus-castus L., MM 6542 


VIOLACEAE 
we lanceolata L.,LB 27859 
iola palmata L., MM 6129; LB 27812; GW 2519 


2542 
Viola pedata L., MM 6138;SJ 16092 
Viola primulifolia L.,SJ 16134;LB 29014;MM 6252: 


SEWP 200 
Viola sororia Willd. var missouriensis (Greene) 
cKinney, LB 28135, 27819; GW 1941, 2443 
Viola villosa Walter, MM 6109;LB 27787;GW 2481, 
2931 


VITACEAE 
Ampelopsis arborea (L.) Koehne, MM 6728, 6728; 


Parthenocissus quinquefolia (L.) Planch., MM 
6126; SJ 16094 
Vitis aestivalis Michx., GW 2580; MM 6558, 6287; 
16274 


Vitis cinerea (Engelm) Millard. var. cinerea, LB 
605, 28132 
Vitis rotundifolia Michx., MM 6124; LB 28120; SJ 
16223 


XYRIDACEAE 

Xyris ambigua Bey. ex Kunth, LB 28715 

Xyris baldwiniana Schult., MM 6580; GW 2711;LB 
30459 


BROWN ET AL., FLORA OF THE BIG SANDY PRESERVE 721 


Xyris ee 5 Chapm. var. curtissii (Malme) Kral, Xyris difformis Chapm. var. difformis, GW 2710 
B 28425; MM 6732 Xyris jupicai L.C. Rich., MM 6888, 6700 


Notes 

Agalinis filifolia— We are following Wunderlin (1998) in merging A.. pulchella 
with A.. filifolia. 

Cerastium pumilum.—Turner et al (2003) mapped this exotic species in six 
northeastern Texas counties with Kaufman County the most southern. Our Polk 
County record and another one from San Jacinto County (both at SBSC) are the 
most southern Texas collections known to us. 

Croptilon hookerianum.—The Polk County collection is the first for the 
pineywoods region of east Texas. The most eastern collection in Turner et al. 
(2003) is from Fayette County. It is distinguished from C. divaricatum and C. 
rigidifolium (the only other Croptilon taxa in Texas) by the lack of non-glan- 
dular trichomes on the distal portions of the stems. 

Diaperia candida.—In the Flora of North America Asteraceae volumes (ined.), 
our Evax taxa are treated in the genus Diaperia. 

Loeflingia squarrosa.—This western taxa appears to be new to Polk County 
(Turner et al. 2003). This, Prunus eracilis,and Croptilon hookerianum are the 
only strictly western sy found on the Big Sandy Creek Unit. These are char- 
acteristic species of xeric sandylands. 

Minuartia muscorum.—For the curious nomen clatural history of this taxon see 
Rabeler (1992). 

Phlox nivalis subsp. texensis.—1T his endemic to southeast Texas anda federally 
and state listed endangered species (Poole & Riskind 1987; Texas Parks and 
Wildlife 2004), was located on the Big Sandy Creek Unit during the study. The 
species has been planted in several locations in the unit by the USS. Park Service, 
but the population located during this study was a naturally occurring one 
which had been known for some years. 

Ranunculus recurvatus.—Our Polk County collection is one of only four coun- 
ties mapped in Turner et al. (2003). 

Silene subciliata.—In addition to Phlox nivalis ssp. texensis, Silene subcilata is 
the only other Texas rare species found on the Big Sandy Creek Unit. This West 
Gulf Coastal Plain endemic is rare in both Texas and Louisiana, but is not fed- 
erally listed. In addition to these two rare West Gulf Coastal Plain endemics, 
the Big Sandy Creek Unit has several other West Gulf Coastal Plain endemics 
(MacRoberts et al. 2002a). These are Baptisia bracteata var. leucophaea, Bapti- 
sia nuttalliana, Berlandiera pumila var. scabrella, Dalea phleoides var. phleoides, 
Echinacea pallida var. sanguinea, Hymenopappus artemistifolius var. 
artemisiifolius, Lobelia puberula, Mimosa hystricina, Oligoneuron nitidum, 
Panicum brachyanthum, Scutellaria cardiophylla, Sisyrinchium sagittiferum, 
Solidago ludoviciana, Tradescantia reverchonii, and Vernonia texana. 


722 BRIT.ORG/SIDA 22(1) 


Sophora affinis—Our Polk County collection is one of only three specimens 
known to us from the pineywoods region in east Texas. The other two collec- 
tions are from the Trinity River bottomlands in Liberty County. 


DISCUSSION 


There are 126 families and 358 genera for the 693 taxa (648 native) on the Big 
Sandy Creek Unit list. Also, 50.9 % of all taxa belong to eight families; they are 
Poaceae (88 taxa), Asteraceae (85 taxa), Cyperaceae (69 taxa), Fabaceae (46 taxa), 
Lamiaceae (19 taxa), Euphorbiaceae (16 taxa), Scrophulariaceae (16 taxa), and 
Fagaceae (15 taxa). The largest genera are Carex (35 taxa), Dichanthelium (19 
taxa), Quercus (13 taxa) Cyperus (11 taxa), and Rhynchospora (11 taxa). 

We compared our list of taxa for Big Sandy Creek Unit with the Polk County 
taxa reported by Turner et al. (2003) in their Atlas of the vascular plants of Texas 
and found that only 70 percent of our taxa are mapped for Polk County in that 
source. This is not surprising considering that until recently little focused col- 
lecting has been undertaken in the Big Thicket region and indicates that basic 
floristic collecting and documentation in the Big Thicket region are still needed. 

Thus, while itis the case that our list isincomplete and that a few taxa may 
no longer exist on the unit, most taxa that have grown in the Big Sandy Creek 
Unit during the past quarter century have been collected (probably 85+ per- 
cent). We estimate that the Big Sandy Creek Unit has about 660 to 750 native 
species/taxa. 


ACKNOWLEDGMENTS 


We thank Guy Nesom (BRIT) and David Riskind (Texas Parks and Wildlife De- 
partment) for their careful review and resulting improvement of this paper and 
the staff of TEX/LL, largely Tom Wendt and Lindsay Woodruff, for facilitating 
the senior authors’s examination of the William Carr specimens. This study 
was supported in part by a National Park Service Cooperative Agreement No. 
CA 14001004 to Paul Harcombe who also provided funds for the page charges. 


REFERENCES 


Autvscl, G. 1979. Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas 
A&M Press, College Station. 

Brown, L.E.,B.R. MacRoserts, M.H.MacRoserts, PA. HAaRcomBe, W.W. Pruess, |S. Ecsik, and D. JOHNSON. 
2005. Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big 
Thicket National Preserve, Tyler and Hardin counties, Texas. Sida 21:1807-2005. 

Desuorets, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. U.S.D.A. Soil 
Conservation Service, College Station, Texas. 

Harcomee, P.A.and P.L. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. 
Unpublished report: U.S. Park Service, Santa Fe, NM. 


BROWN ET AL., FLORA OF PRESERVE 723 


Harcomee, PA., J.S. Guzenstein, R.G. Knox, $.L. Orzett, and E.L. Brioces. 1993. Vegetation of the 
longleaf pine region of the West Gulf Coastal Plain. Proc. Ann. Tall Timbers Fire Ecol. 
Conf. 18:83-103. 

Jones, S.D., J.K. Wipre, and P.M. Montcomery. 1997. Vascular plants of Texas. Univ. Texas Press, 
Austin. 

Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. First edition. |n:J.T.Kartesz and 
C.A.Meacham. Synthesis of North American flora.Version 1.0.North Carolina Botanical 
Garden. Chapel Hill. 

MacRoserts, M.H., B.R. MacRoserts, B.A. Sorric, and R.E. Evans. 2002a. Endemism in the West 
Gulf Coastal Plan: importance of xeric habitats. Sida 20:767-780. 

MacRoserts, B.R., M.H. MacRoseets, and L.E. Brown. 2002b. Annotated checklist of the vascu- 
lar flora of the Hickory Creek Unit of the Big Thicket National Preserve, Tyler County, 
Texas. Sida 20:781-795. 

Marks, PL. and PA. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas. 
Ecol. Monogr. 51:287-305. 

McLeop, C.A. 1971. The Big Thicket forest of east Texas. Texas J. Sci. 23:221-233. 

Parks, H.B. and V.L. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas 
Agricultural Experiment Station, College Station. 

Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A&M. Press, College 
Station. 

Poote, JM. and DH. Riskino. 1987. Endangered, threatened, or protected native plants of 
Texas. Texas Parks and Wildlife Dept., Austin. 

TEXAS PARKS AND WILDLIFE & THE NATURE CONSERVANCY, AusTIN, Texas. 2004. List of the rare plants of 
Texas. http://nature.org/whwerwework/northamerica/states/texas/files/ 
listofrareplants/pdf. 

RaBeLer, R.K. 1992. A new combination in Minuartia (Caryophyllaceae). Sida 15:95—96. 

Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. 
Sida Bot. Misc. 24: 1-888. 

Watson, G.E. 1979. Big Thicket plant ecology: an introduction. Big Thicket Mus. Publ. Ser, 
No. 5, Saratoga, Texas. 

Watson, G.E. 1982. Vegetational survey of Big Thicket National Preserve. Unpublished re- 
port. Big Thicket National Preserve, Beaumont, TX. 

WunNbeERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida. 
Tampa. 


724 BRIT.ORG/SIDA 22(1) 


THE NEW YORK BOTANICAL GARDEN PRESS 


Flora Genérica de los Paramos 
Guia Ilustrada de las Plantas Vasculares 


Petr Sklendr, James L. Luteyn, Carmen Ulloa Ulloa, Peter M. Jorgensen and Michael O. Dillon 


Memoirs of The New York Botanical Garden 
Volume 92 

ISBN-0-89327-468-2 

Order no. MEM 9 

$85.00 


520 pages 
n Spanish with an English abstract 


=a 


P4ramo is one of the richest nce elevation mountain ecosystems in the world with high biological 
diversity and endemism, but it is ecologically fragile. This book provides keys, desc a era distrib- 
utional information, and illustrations for the dence ation of the 127 families and 540 genera of 


vascular plan 


le | i s de alta montafa mas rico del Mundo con una diversidad 


es 


ramo es 
bioldgica y un endemismo altos, pero es frégil ecolégicamente. Este libro proporciona claves, 


descripciones, informacién acerca de la distribucién e ilustraciones para la identificacién de 


las 127 familias y 540 géneros de plantas vasculares 


“three-digit r Manca urd nani on ont 
: ofA rie Ez Ord ath P sed withour this securi 


: instirudon, 
Se Tee ee ie eee eee 


SIDA 22(1): 724. 2006 


THE NORTH-SOUTH TRANSITION OF FLORA 
ACROSS ARKANSAS: A PRELIMINARY 
PHY TOGEOGRAPHICAL ANALYSIS 


Michael H. MacRoberts and Barbara R. MacRoberts 


Bog Research 
740 Columbia 
Shreveport, Louisiana 71104, U.S.A. 
and 


Herbarium, Museum of Life Sciences 
Louisiana State University in Shreveport 
Shreveport, Louisiana 71115, U.S.A. 


— 


ABSTRACT 


1 ] Ae A : hi i | 1 Using quan- 


titative nethods we examined the north- south f floristic feaneiaga ACTOSS Denese to see if phyto- 
geographic regions are detectable by abrupt changes in the flora. We found only gradual change 
even though there are abrupt and major physiographical discontinuities. 


RESUMEN 


Los BleeeOssaes han dividido Arkansas en provincias fisiogeograficas y ecorregiones. Usando 
uantitativos, examinamos las transiciones norte-sur a través de Arkansas para ver si las 


regiones fitogeograficas son  detectabes por Cambios peeEuDios en la flora. Hemos encontrado solo 
y mayores. 


}: 


cambio gradual aunque hay 


3 


INTRODUCTION 
Virtually all ¢ hic and ecoregional mappers treat the Interior High- 
lands of Arkansas, Illinois, Kansas, Missouri, and Oklahoma as being distinct 
from surrounding areas. While most appear to agree on a physiogeographic 
map, they present conflicting interpretations of the biotic provinces and 
ecoregions (e.g., Dice 1943; Braun 1950; Gleason & Cronquist 1964, Kuchler 1964; 
Foti 1974; Pell 1983; Omernik 1986; Takhtajan 1986; Thorne 1993; Bailey et al. 
1994; Keyset al. 1995; Weakley et al. 1998; Foti & Bukenhofer 1998; Ricketts 1999). 
For example, Dice (1943), Takhtajan (1986), and Thorne (1993) consider the In- 
terior Highlands and Gulf Coastal Plain to be different provinces, Bailey et al. 
(1994) and Keys et al. (1995) break the Interior Highlands into two provinces, 
Omernik (1986) divides the area into six ecoregions, and Weakley et al. (1998) 
divide it into four ecoregions. 

While it is unquestionably the case that the Ouachita Mountains and the 
Ozark Plateau are physiogeographically distinct from surrounding areas, no- 
tably the West Gulf Coastal Plain to the south, it is not so certain that the area is 
particularly distinct floristically. Zollner et al. (2005:1788), speaking of the In- 


SIDA 22(1): 725 — 734. 2006 


726 BRIT.ORG/SIDA 22(1) 


terior Highlands in general, question its uniqueness: “for all of its physiographic 
uniqueness, including age, long-term isolation from its moiety—the Appala- 
chian region—, and its reputation as a ‘well-known refugium’ (Meyer 1997:364) 
the Interior Highlands] shows surprisingly little floristic uniquity from sur- 
rounding regions.” This was also recognized years ago by Palmer (1921) when 
he pointed out that about 90 percent of the woody flora of the Ozark region 
also occurs in the Gulf Coastal Plain. 

This point was brought home to us forcefully when we conducted a floris- 
tic inventory of the Ouachita Mountains Biological Station in Polk County, Ar- 
kansas (MacRoberts et al. 2005). Having extensive experience with the flora of 
the West Gulf Coastal Plain, but none with the Ouachita Mountains, we ex- 
pected, since the physiogeography was so different and since biogeographers 
have separated the Interior Highlands from the Coastal Plain, that the two ar- 
eas would be quite floristically distinct. Surprisingly they are not. The flora of 
the Ouachita Mountains Biological Station presented some species unknown 
to us, but not many. 

Ina previous paper, we documented the east-west floristic transition across 
central Texas in which the eastern flora precipitously drops out and is replaced 
by a western flora and vice versa (MacRoberts & MacRoberts 2003). In this 
study, using similar methods we examine the north-south floristic transition 
across Arkansas, i.e., from the West Gulf Coastal Plain in the south into the In- 
terior Highlands in the north (Fig. 1). Western Louisiana and southern Arkan- 
sas are physiographically the Gulf Coastal Plain (Fenneman 1938; Brouillet & 
Whetstone 1993) and the Atlantic and Gulf Coastal Plain floristic province 
(Takhtajan 1986; Thorne 1993). The Interior Highlands, on the other hand, are 
part of the Ozark and Ouachita physiographic region (Fenneman 1938; Brouillet 
S Whetstone 1993) and the Appalachian floristic province (Takhtajan 1986; 
Thorne 1993). 

The purpose of this paper is to provide data on 1) how different floristi- 
cally the Interior Highlands is from the West Gulf Coastal Plain, 2) the effect of 
the physiogeographic differences on the north-south floristic transition of flora 
from the Coastal Plain into the Interior Highlands, and 3) the floristic relation- 
ship between the Interior Highlands and the regions that surround it. 


— 


— 


STUDY SITES 


The Interior Highlands and the Gulf Coastal Plain have been described in numer- 
ous publications (Fenneman 1938; Foti 1974; Pell 1983: Bryant et al. 1993; Skeen 
et al. 1993; Dale & Ware 1999; Delcourt & Delcourt 2000) and will not be redescribed 
here except to say that the former region consists of moderate elevation moun- 
tains of Paleozoic sedimentary rock with numerous swift clear streams and riv- 
ers and both deciduous and pine-hardwood forests; the latter region consists of 
low to gently rolling topography of Cretaceous and Tertiary silts and clays char- 


NEGCeae / 
mae” 2 


Ouachita ae ae nu 


Reece 
ors, 


3 


or 


ae 


A 


Fic. 1. Location of Interior Highlands and Gulf ¢ | 
and Baxter (B), Garland (G), Newton (N), and Polk (P) counties. 


728 BRIT.ORG/SIDA 22(1 


=> 


acterized by forests of pine and hardwoods, with swamps, and sluggish muddy 
streams and rivers, and oxbow lakes. Precipitation is about the same in both 
areas, with temperatures slightly cooler in the Interior Highlands. 


METHODS 


1) As we pointed out in the introduction, we have recently conducted a floristic 
survey of the 211 hectare Ouachita Mountains Biological Station in Polk County, 
Arkansas (MacRoberts et al. 2005). It is located 174 km north of Louisiana in 
the Ouachita Mountains and has an elevation range of between 395 and 622 
meters. It is dominated by pine-hardwood forests on shale, chert, novaculite, 
siltstone, and sandstone. It could not be more different edaphically and topo- 
graphically from the West Gulf Coastal Plain. We have documented 315 native 
species on the station. We compared this list to floristic lists for Louisiana (Tho- 
mas & Allen 1993-1998; Kartesz & Meacham 1999) to determine the percent- 
age of species in common. While the Ouachita Mountains Biological Station is 
a very small area given the size and diversity of the Ouachita Mountains, it pro- 
vided a first comparison of the flora between the West Gulf Coastal Plain and 
the Ouachita Mountains. 

2) In order to obtain a more detailed understanding of the possible effect 
of the Interior Highlands on plant distribution, using the Thomas and Allen 
(1993-1998) atlas we randomly selected 254 native species occurring in north- 
ern Louisiana (north of TIN) that also occur in Arkansas (Smith 1988, 1994). 
Using Kartesz and Meacham (1999), we determined how many of these occur 
in Missouri, approximately 380 km to the north. The sample size is only about 
10 percent of the native species but sufficient to compare the flora between the 
regions. 

3) As an independent test of Method 2, using Hyatt (1993) we determined 
the percentage of native species occurring in Baxter County, Arkansas, on the 
Missouri border in the Ozark Plateau, that also occur in the Gulf Coastal Plain 
of Texas and Louisiana (Thomas & Allen 1993-1998: Kartesz &@ Meacham 1999; 
Turner et al. 2003). Naturally, some of the species in this sample are the same as 
those in Method 2. 

4) Using the data in Method 2, we plotted the Arkansas county distribu- 
tion of the 254 native species. Because all counties are not collected equally 
(e.g., Jackson County has 275 reported taxa while Washington County has 1,301 
reported taxa), we expressed the results as percentages; that is, we divided the 
number of species in our sample reported for each county by the total species 
reported for each county. Thus, of the 254 species in the sample, 99 are reported 
for Miller County, and Smith (1988) reports 544 species for Miller County (99/ 
544 = 18%). 

5) Having found that Method 4 did not reveal sharp distributional 
discontinuities in the flora across Arkansas (see “Results”), using Kartesz and 


, BIOGEOGRAPHICAL ANALYSIS OF THE ARKANSAS FLORA 729 


Meacham (1999) we surveyed the entire flora of North America and found all 
of the native species that were reported to occur in Missouri and Arkansas (but 
not in Louisiana) and in Louisiana and Arkansas (but not in Missouri) irrespec- 
tive of where else they occurred. Using Smith (1988), we then plotted these spe- 
cies by Arkansas counties according to whether they were Missouri/ Arkansas 
species or Louisiana/ Arkansas species (i.e, coming from the north or the south). 
Our sample consisted of 499 species: 272 Missouri/Arkansas and 227 Louisi- 
ana/ Arkansas. Our Buea es) procedure was the same as in Method 4. Our hope 
was to magnify the curve b inating species that occurred in all three states. 

6) In order to determine the overall North American floristic association 
of the Interior Highlands, using Smith (1988) we randomly selected 296 native 
species from Garland County, Arkansas, which is entirely within the Ouachita 
Mountain region, and 293 native species from Newton County, Arkansas, which 
is entirely within the Ozark Plateau region (both counties being well collected) 
and, using Kartesz and Meacham (1999), plotted their state/regional occurrence 
expressed as percentage of species in common. 

All of these measures are of presence/absence, not abundance. The data 
necessary to investigate abundance are not yet available on a large scale. Thus, 
we recognize that some taxa may be widely distributed across the region while 
others may not. Also, while we recognize the limitation of the main data sources 
for this paper, we have made every effort to adjust to those limitations. But since 
the study is a preliminary analysis, it can absorb some error without compro- 
mising the general conclusions. The paper provides a problem and an analysis 
that should be followed up by a more extensive sample of species and more 
points along the continuum. 


RESULTS 


1) Of the 315 species on our Ouachita Mountain Biological Station list, 287 or 
91% occur in Louisiana entirely within the West Gulf Coastal Plain. 

2) Of the 254 species in our north Louisiana sample, 209 or 82% reached 
Missouri and 45 or 18% stopped in Arkansas. 

3) Eighty-two percent of the native species that occur in Baxter County on 
the Missouri border also occur in the West Gulf Coastal Plain of Louisiana and 
Texas. 

4) Figure 2 summarizes the results of Method 4. Counties on the southern 
tier of Arkansas averaged around 18% and 19% of the sample, and counties on 
the northern tier averaged between 12% and 16% of the sample. No marked or 
abrupt shifts in flora were detected that correlated with the natural 
physiogeographic divisions of Arkansas. 

5) Figure 3 summarizes the results of Method 5. No marked or abrupt shifts 
in flora were detected that correlated with the natural physiogeographic divi- 
sions of Arkansas. 


730 BRIT.ORG/SIDA 22(1) 


* 12 [12] 13] 46 Lali eaey" 

© 12 12 . 

4 ere a 
je tial fel 


"18! 


f ag 


oe 


Pen ae ae 


Fic. 2. P t by £7CA] 


L ] . \ 
y t \ TUPLE CAPIdHaUlOit). 


6) Figure + shows that the Ouachita Mountains has about 91% of its flora 
in common with the West Gulf Coastal Plain and the Ozark Plateau has about 
82% of its flora in common with the West Gulf Coastal Plain. It also shows that 
the Ouachita and Ozark mountain/plateau regions are not only continuous flo- 
ristically with the surrounding regions but virtually indistinguishable from 
them. 


Laer | 
— 


DISCUSSION 


As noted in the introduction, biogeographers have presented a number of 
schemes for subdividing the Interior Highlands. These range from putting the 
Interior Highlands and Gulf Coastal Plain into separate biotic provinces or 
ecoregions to dividing the Interior Highlands into two provinces, and so on. 


MACROBERTS AND MACROBERTS, BIOGEOGRAPHICAL ANALYSIS OF THE ARKANSAS FLORA 731 


Pee 2 | I 
TUPUTTET CAPId- 


Our analysis shows that the vast majority of plants occurring in north Louisi- 
ana and east Texas (West Gulf Coastal Plain) are also found in central and north- 
ern Arkansas. There is a gradual transition of species from the West Gulf Coastal 
Plain into the Interior Highlands and vice versa, and there are no marked or 
abrupt shifts correlated with the natural divisions of Arkansas, indicating that 
there are no major physiographical barriers. The only place we may have de- 
tected a possibly sharper transition is between the Ozarks and the Mississippi 
floodplain, but even here the transition is not precipitous. 

The Interior Highlands are not floristically distinct from adjacent regions. 
The Ouachita flora as represented by Garland County, Arkansas, is closely re- 
lated to the flora of all surrounding states especially Oklahoma, Louisiana, 
Mississippi, Tennessee, Kentucky, Alabama, Georgia, and the Carolinas (Fig. 4). 
The Ozark flora as represented by Newton County, Arkansas, is very similar to 
the Ouachita flora but, as would be expected, has a more northern affiliation. 
The Ozark region has its strongest floristic affinity with Oklahoma, Missouri, 
Illinois, Indiana, Kentucky, Tennessee, Ohio, Mississippi, Alabama, and Geor- 
gia (Fig. 4). If any generalization is necessary, the Interior Highlands as Zollner 
et al. (2005: 1788) have pointed out “shows surprisingly little floristic uniquity 
from surrounding regions.” 

Perhaps some or most of this lack of uniqueness is the result of movernents 
of species north-south for the last two million years of 20 glaciations, with strag- 
glers both north and south. This pattern of plant movement has been docu- 
mented extensively, notably by pollen analysis but also by fossil plants (Givens 
& Givens 1987, Delcourt & Delcourt 2000). Also, edaphic conditions are not 
entirely dissimilar along this north-south continuum, with high rainfall and 
generally mild climates. 


732 BRIT.ORG/SIDA 22(1) 


. flnrice: Lins? fh) la (lnft) 1 far] Wa {eiahe) Ct 
1c.4.N +h A 


of species in common. 


What these results mean for biogeographic classification and mapping is 
not entirely clear. Takhtajan (1986) has defined “province” differences as involv- 
ing low levels of generic endemism and high levels of species endemism. 
“Ecoregion” is generally defined as a large area consisting of a distinct assem- 
blage of natural communities and species with endemism being fairly high at 
the species and subspecies level (MacRoberts & MacRoberts 2004). The West 
Gulf Coastal Plain and the Atlantic and Gulf Coastal Plain have been the sub- 
ject of recent endemic studies (Sorrie @ Weakley 2001; MacRoberts et al. 2002). 
The West Gulf Coastal Plain has approximately 100 endemic taxa, three at the 
generic level. The Interior Highlands has only about 37 endemics, none at the 
generic level (Zollner et al. 2005). Setting aside the fact that no precise “index” 
has been proposed to test for provincial or ecoregional status, it seems from our 
analysis that the Interior Highlands and Gulf Coastal Plain are too similar flo- 
ristically to be classified as different provinces or ecoregions. Perhaps some lower 
chorionomic category might better classify these two areas. 


ACKNOWLEDGMENTS 


Douglas Zollner, The Nature Conservancy, commented on an earlier version of 
this paper. George P. Johnson and an anonymous referee commented on the paper. 


REFERENCES 


Baivey, R.G., PE. Avers, T. Kine, and W.H. McNas. (eds.). 1994. Ecoregions and subregions of the 
United States. (2 maps). U.S. Geological Survey, Washington, D.C 

Braun, E.L. 1950. Deciduous forests of eastern North America. Hafner, New York. 

Brouittet, L.and R.D. Wuerstone. 1993.Climate and Physiography.|n: Flora of North America 
Editorial Committee, eds. Fl. North Amer.1:15—46. Oxford Univ. Press, New York. 

Bryant, W.S., W.C. McCome, and J.S. FrauisH. 1993. Oak-Hickory forests. In: W.H. Martin, S.G. 


MACROBERTS AND MACROBERTS, BIOGEOGRAPHICAL ANALYSIS OF THE ARKANSAS FLORA 733 


Boyce, and E. Echternacht, eds. Biodiversity of the southeastern United States: upland 
terrestrial communities. John Wiley, New York. Pp. 143-201. 

Date, E.E. and S. Ware. 1999. Analysis of oak-hickory-pine forests of Hot Springs National 
Park in the Ouachita Mountains, Arkansas. Castanea 64:163-174. 

Detcourt, H.R.and PA. Detcourr. 2000. Eastern deciduous forests. In: M.G. Barbour and W.D. 
Billings, eds. North American terrestrial vegetation. Cambridge Univ.ersity Press, New 
York. Pp. 357-395. 

Dice, L.R. 1943. The biotic provinces of North America. Univ. Michigan Press, Ann Arbor. 

FeNNeMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill, New York. 

Foti, T.L. 1974. Natural divisions of Arkansas. In: Arkansas natural areas plan. Arkansas De- 
partment of Plannning, Little Rock. Pp. 11-37. 

Foti, T.L.and G.A. BukenHorer. 1998. A description of the sections and subsections of the 
Interior Highlands of Arkansas and Oklahoma. J. Arkansas Acad. Sci. 52:53-62. 

Givens, C.R.and F.M. Givens. 1987. Age and significance of fossil white spruce (Picea glauca), 
Tunica Hills, Louisiana-Mississippi Quaternary Quart. 27:283-296. 

Gieason, H.A.and A.Cronauist 1964. The natural geography of plants.Columbia Univ. Press, 
New York. 

Hyatt, PE. 1993.A survey of the vascular flora of Baxter County, Arkansas. Castanea 58:115- 


Kartesz, J.T.and C.A. MeacHam. 1999. Synthesis of North American flora. Version 1.0. North 
Carolina Botanical Garden, Chapel Hill. 

Keys, J.E., CA. CARPENTER, S. Hoos, F. Koenic, W.H. McNas, W.E. Russet, and M. SmitH. 1995. Eco- 
logical units of eastern United States—first approximation. U.S.D.A. Forest Service, At- 
lanta, GA. 

KucHter, A.W. 1964. Potential natural vegetation of the conterminous United States. Amer. 
Geogr. Soc., Special Publ. 36. New York. 

MacRoserts, M.H., B.R. MacRoserts, B.A. Sorrig, and R.E. Evans. 2002. Endemism in the West 
Gulf Coastal Plain:importance of xeric habitats. Sida 20:767-780. 

MacRoserts, M.H. and B.R. MacRoserts. 2003. The east-west transition of flora in Texas: a 
biogeographical analysis. Sida 20:1693-1 700. 

MacRoserts, M.H. and B.R. MacRoserts. 2004. The post oak savanna ecoregion: a floristic 
assessment of its uniqueness. Sida 21:399-407. 

MacRoseets, B.R, M.H. MacRoserts, and T.D. Marsico. 2005. Preliminary survey of the vascular 
flora of the Ouachita Mountains Biological Station, Polk County, Arkansas. Bull. Mus. 
Life Sci. 13:1-19. 

Meyer, F.G. 1997. Hamamelidaceae. In: Flora of North America Editorial Committee, eds. 
Flora of North America, volume 3. Oxford University Press, New York. Pp. 362-367. 

Omernik, J.M.1986.Ecoregions of the conterminous United States. Suppl. Ann. Assoc. Amer. 
Geogr. 77 (1).Map. 

Pavmer, E.J. 1921.The forest flora of the Ozark region. J. Arnold Arbor. 2:216-232. 

Pet, W.F. 1983. The natural divisions of Arkansas:a revised classifications and description. 
Natural Areas J. 3:12-23. 


734 BRIT.ORG/SIDA 22(1) 


Ricketts, T.H. 1999. Terrestrial ecoregions of North America. Island Press, Washington, D.C 

Skeen, J.N., PD. Doerr, and D.H. Van Lear. 1993. Oak-hickory-pine forests. In: W.H. Martin, S.G. 
Boyce, and E. Echternacht, eds. Biodiversity of the southeastern United States: upland 
terrestrial Communities. John Wiley, New York. Pp. 1-33. 

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2nd ed. 
Privately Printed, Fayetteville, AR. 

Sori, B.A.and A.S.Weaktey.2001.Coastal plain vascular plant endemics: phytogeographic 
patterns. Castanea 66:50-82. 

TAKHTAJAN, A. 1986. Floristic regions of the World. University of California Press, Berkeley. 

THomas,R.D.and C.M. ALLEN. 1993-1998. Atlas of the vascular flora of Louisiana. The Nature 
Conservancy, Baton Rouge. 3 Vols. 

THorne, R.F. 1993. Phytogeography. In: Flora of North America Editorial Committee, eds. 
Flora of North America, volume1. Oxford University Press, New York. Pp. 132-153. 

Turner, B.L., H. NicHots, G. Denny, and O. Doron. Atlas of the vascular plants of Texas. Sida Bot. 
Misc. 24:1-888. 

Weak ey, A.S.,K.D. Patterson, S.LANDALL, and M. Pyne. 1998. International classification of eco- 
logical Communities: terrestrial vegetation of the southeastern United States. The Na- 


ture Conservancy, Chapel Hill. 
Zouuner, D., M.H. MacRoserts, B.R. MacRoserts, and D. Lapp. 2005. Endemic vascular plants of 
the Interior Highlands, U.S.A. Sida 21:1781-1791. 


HIGH LEVELS OF SEED INVIABILITY AMONG 
SEVEN POPULATIONS OF THE ENDANGERED 
SHORTS GOLDENROD (SOLIDAGO SHORTII ASTERACEAE) 


Patrick J.Calie Robert F.C. Naczi 


Dept. of Life Sciences The Claude Phillips Herbarium 
235 Mone selene Bldg. Delaware State University 
Eastern Kentucky University Dover, Delaware 19901, U.S.A. 
Biemend: Kentucky 40475, U.S.A. 
pat.calie@eku.edu 
Christina Shackleford Jessica Caicedo 
Dept. orenvionmenta! Health Science ries one Sciences 
Eastern ea Vary East y University 
Richmond, Kent 40475, U.S.A, eae Kentucky 40475, U.S.A, 
ABSTRACT 


The physical conditions of mature seeds If x Kentucky popu- 


lations and one Indiana population of the endangered species ‘Short’s eaieeed. (Solidago shortii, 
Asteraceae) A sadn ee bsenes oe eae Reutheny populations levealed the majority (71%- 


OOO | 
6) 


I 1 insect preda- 
tor or to unsuccessful fertilization or aie ryo tion. Successful seedling ee from phe- 
notypically normal seeds eet rom 71% to 96% in the ed Kentucky populations. The majority 

of the damaged seeds were necrotic, ranging from 62%- 90%, The Indiana popwlanon exiubited the 
lowest ee of herbivory ee) ebbeived for the ESS ae congener gold anadensis 


‘5 


1andS 


or adjacent to thre lati f Short’s 
ae exhibited lower levels of necrosis Ae higher eee of herbivory. A ee congener species 
(S. gigantea) exhibited equal levels of the two seed conditions (herbivory and necrosis). The influ- 
ence of seed inviability upon the persistence of natural populations of Short’s goldenrod is at present 
een 


RESUMEN 


dici Ficicg | mil] 


maduras de 2 ope amenazada “Short’s goldenrod” (Solidago 
shortii, A yi inad i 1 


Kentucky y una poblacion de Indiana. 
La ayer a Hae sepals de poblaci las al azar en ERentiesy Pisce arOne un numero 
reducido ] herbivoria, infertilidad 
estas ebenvabiones Semillas normales encontradas en las poblaciones de Kent oe dene ron 
plantulas que se desarrollaron con éxito en un 71% a 96%. En la mayoria de las semillas danadas los 
niveles de necrosis fluctuaron en un 62%-90%. La poblacion de Indiana presenté el menor nivel de 
herbivoria (2%) ob vado ena epee En dos Spee: de “goldenrod” (S. canadensis var. altissima y 
S lis) distribuid | “Short’s goldenrod” se observaron bajos 
ivelesd isyal iveles de herbivoria. E le “goldenrod” (S. gigantea) 
se ena Sonetone similares de mopivens y necrosis. Se deseoncte is insluencte de 
ae 


a persl 


SIDA 22(1): 735 — 748. 2006 


736 BRIT.ORG/SIDA 22(1) 


INTRODUCTION 


Short’s goldenrod (Solidago shortii, Asteraceae) isan endangered species extant 
in the Blue Licks area of north central Kentucky and in one locality in Indiana 
(Homoya & Abrell 2005). A history of the discovery of this species, a descrip- 
tion of the Kentucky localities in which this plant is currently found, and gen- 
eral features of the species’ ecological life history are summarized in Baskin et 
al. (2000). 

As with many endangered species, a key question remains unanswered: 
Why does this species have such a limited geographical distribution? In an ef- 
fort to answer this question, an extensive body of knowledge has emerged re- 
garding the autecology of Short’s goldenrod (e.g. Buchele et al. 1989), involving 
investigations of both abiotic and biotic factors and their possible impact on 
the occurrence of this rare species. A later series of investigations implicated 
interspecific competition, primarily from introduced species, as being a lead- 
ing candidate greatly interfering with establishment of Short’s goldenrod seed- 
lings (Walck et al. 1999a, 1999b). 

One possible biotic component contributing to the rar 2 of the species is 
its reproductive biology, e.g. successful production of viable offspring. The first 
studies focusing on S. shortii seed germination were conducted by Buchele et 
al. (1991), that were later greatly expanded by Walck et al. 1997a, 1997b, 1997c, 
1997d, 1997e, 1997f, 1999c). The primary (but not sole) emphasis in these stud- 
ies was to determine the biotic and abiotic factors most responsible for success- 
ful seed germination and seedling survival and the range of conditions (e.g. light, 
temperature, moisture, cold stratification, etc.) required for optimum germina- 
tion. In the first set of germination experiments (Buchele et al. 1991) seeds were 
obtained from two populations of Short’s goldenrod; in the second set (Walck 
et al. 1997a) one population was sampled. 

In our investigations involving the biology of Short’s goldenrod, we noticed 
high levels of seed damage in individuals from several Kentucky populations 
and asked whether differences existed among populations in reproductive po- 
tential, ie. the production of phenotypically normal seeds and developmen- 
tally normal seedlings. We posed the following questions regarding the repro- 
ductive biology of S. shortii. 1) What types of seed damage are present? What 
types of variation, if any, exists at the population level? Are some populations 
more prone to particular damages than others? 2) What quantitative variation, 
if any, exists both within and among selected Kentucky populations of Short’s 
goldenrod in the levels of different types of seed damage and the production of 
normal seedlings? 3) Do the common congeners of S. shortii in the Blue Licks 
vicinity (ie. S. canadensis var. altissima and S. nemoralis) and at the Indiana 
site (S. gigantea and S. nemoralis) exhibit similar levels of normal seed produc- 
tion as S. shortii? Are there statistically significant differences between the Ken- 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 737 


tucky and Indiana populations of Short’s goldenrod with respect to normal seed 
production? 


MATERIALS AND METHODS 


To address the first two questions, matur 1s (cypselae) of S. shortii were col- 
lected in mid-December 2001 from six Kentucky populations designated as #1, 
2,5, 7,8, and 12 (Evans 1987; Smith et al. 2004). This time was chosen to ensure 
the collection of mature seeds, as noted in earlier studies (Buchele et al. 1991a; 
Walck et al. 1997a). The sampled populations are found within a 2.2-km area of 
Blue Licks Battlefield State Park in northern Kentucky, and were chosen due to 
the prevalence of flowering stems and physical accessibility. Ten individual 
stems were non-randomly sampled from each population as follows. The ini- 
tial sampled plant stem was a minimum of 10 meters from the eg ae of 
that specific population (based on the GIS ing data from Smith et al. 2004). 
Due tothe non-random distribution of these plants, the cluster of plants in clos- 
est proximity to the first sampled stem was then sampled, this process being 
repeated until ten plants were sampled. To maximize the probability of sam- 
pling separate genets, each sampled stem was separated from any neighboring 
sampled stems by at least 3 meters. Portions of mature infructescences were 
removed from the stems, placed in a paper coin envelope, and stored at 4°C in 
the dark for 12-16 weeks. 

Following cold storage, the dev tal fate of individual seeds sampled 
from the Kentucky populations in 2001 was followed by the use of 96-well 
microtiter plates (Fig. 1) as germination chambers prepared as follows. A mini- 
mal mineral salts medium (Hickok & Warne 1998) containing 1% agar was au- 
toclaved and then aliquotted into each well (200 ml of medium/well) with a 
multichannel pipettor, then allowed to cool for 24 hours. Seeds were harvested 
by first scraping along the infructescence with forceps to dislodge the fruits 
and then collecting the seeds into a Petri plate. Single seeds were removed (with- 
out regard for condition) by grasping the fruit’s pappus with forceps until the 
appropriate number of seeds was obtained. Each seed (in the majority of cases 
24 seeds per plant, with exceptions noted below) was then carefully imbedded 
half its length into the solidified agar within a single well, with the pappus 
extruding above the medium. The plate cover was labeled with the location and 
identity of each seed source, secured with parafilm, and the plates incubated 
under the following conditions. A daily photoperiod of 14 hours was provided 
by a bank of cool white fluorescent lights (2OW), that produced a photosyn- 
thetic photon flux (400-700 nm) at the germination plate level of approxi- 
mately 20-25 umol m™ sec! The dark period temperature was 20°C and the 
light period temperature was 25°C. A seedling was scored as exhibiting posi- 
tive Viable Seedling Development after emergence of the radicle and a mor- 


] 


738 BRIT.ORG/SIDA 22(1) 


WiahIAc 1: n | (MICH 


example, in column 8 well # (in d di ler) 2,6-10 all contain positive VSD individuals 


phologically normal set of cotyledons that persisted for at least one week post- 
emergence. All plates were incubated for 14 days. 

Preliminary experiments.— Two preliminary experiments were 
to address several experimental parameters. To determine if the damaged seeds 
were capable of either germination or forming normal seedlings, both necrotic 
aiid herbivore-damaged seeds sampled from 10 plants from each of the six popu- 
lations (n=379) were incubated in germination medium for 14 days. In the sec- 
ond experiment the effect of cold-storage on seed germination was evaluated 
with a sample of phenotypically normal seeds (5 populations, 24 seeds/popu- 
lation) incubated in germination medium. Due to an insufficient number of 
normal seeds, data from population #12 were not included. 

Intraspecific comparisons.—Out first experiment of this type involved the 
incubation of a random sample of seeds regardless of physical condition 
(n=1,440) from each of six populations in culture medium, to determine the 
level of viable seedling development (VSD) from a general sample. The second 


ondiicted 
Cd 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 739 


analysis was conducted to determine the actual levels of normal vs. damaged 
seeds in a sample (n=1,199) from all six populations. For each population, 10 
plants were sampled, 20 seeds per plant. For population #2, the seed sample 
size was n = 199. To address the issue of the level of seed damage due to either 
herbivory or necrosis, seeds from five populations were examined for physical 
condition (n=2,431). Ten plants from each population were sampled, typically 
50 seeds per plant, the number of seeds varying in some cases due to availabil- 
ity. Due to an insufficient number of seeds, data from population #8 were not 
included. For the statistical analysis, sample size was reduced to 9 plants per 
population, due to the necessity of having equal sample size per population (all 
sampled populations had at least 9 plants with 50 seeds/plant, but not all popu- 
lations had 10 plants with 50 seeds/plant). 

Interspecific comparisons.—To address the third question mature 
infructescences were collected in mid-December 2003 in the Blue Licks vicin- 
ity from 10 plants of S. shortii and 10 plants of S. canadensis var. altissima from 
population #5, and from 10 plants of S. shortii, 5 plants of S. nemoralis and 5 
plants of S. canadensis var. altissima from population #1. Seeds were collected 
from 10 plants of the Indiana population of Short’s goldenrod and from 5 neigh- 
boring plants of S. gigantea and S. nemoralis each in mid-December 2004. For 
each examined plant, 20 seeds were sampled. Seeds from this final phase of the 
investigation were examined for their physical condition alone. For the Ken- 
tucky populations of S. shortii data from two different collecting seasons (2001 
and 2003) are pooled as no statistically significant difference exists between 
the two seasons (t test; population # 1: t = 1.8, p =0.081; population #2: t = 0.33, p 
= (0.75). For the S. canadensis population data, two different populations (#1 and 
#5) are pooled as no statistically significant inter populational difference exists 
(t test, t= 18, p= 0.12). 

Statistical Analyses.—A|l statistical analyses were conducted using the 
software program SYSTAT version 11 (SYSTAT Software, Inc.). To test hypoth- 
eses of no statistically significant difference in values of the mean between or 
among groups, we used the t-test and analysis of variance (ANOVA), respec- 
tively, except in those cases when assumptions of these tests were violated. 
When the data were not normally distributed and/or group variances were het- 
erogeneous, we used the Mann-Whitney Test or Kruskall-Wallis Test, which 
are nonparametric analogs of the t-Test and ANOVA, respectively (Zar, 1999). 


RESULTS 


Preliminary experiments.—In the first experiment in which damaged and ne- 
crotic seeds were incubated in germination medium, all were inviable as deter- 
mined by the absence of either a radicle or a set of cotyledons. The second ex- 
periment using phenotypically normal seeds was conducted to determine the 
percentages of VSD using the cold storage treatment prior to incubation in seed 


740 BRIT.ORG/SIDA 22(1) 


germination medium. The VSD values ranged from 71% (population #2) to 96% 
(population #8) (Table 1. 

Intraspecific comparisons.—A preliminary visual examination of the Ken- 
tucky S. shortii seeds revealed two distinct morphological classes. Phenotypi- 
cally “normal” seeds had a buff to tan color, were ellipsoid in shape (Fig. 2A), 
and several examined individuals contained fully developed embryos as deter- 
mined by visual examination. The second class, consisting of physically dam- 
aged seeds, were of two types. One type exhibited either small holes bored into 
the seed/fruit coat (Fig. 2B) or a broken end. We term this group as being ex- 
posed to “herbivory,” since the damage appears to be insect-induced. The sec- 
ond group, termed “necrotic,” consisted of seeds that were either very dark brown 
to black, or were of typical color but deflated (Fig. 2C). An examination of sev- 
eral seeds of the former type revealed a shrunken, black mass assumed to be the 
remnants of an embryo. 

Our first experiment of this type involved the incubation of a random 
sample of seeds regardless of physical condition (n=1,440) from each of six 
populations in culture medium, to determine the level of viable seedling devel- 
opment (VSD) from a general sample (Table 2). The levels of VSD were rather 
low ranging from 3% for population #12 to 33% for population #8. The differ- 
ences in levels of VSD between populations are not significant (ANOVA, F = 2.0, 
p= 0.097). 

To determine the actual levels of normal vs. damaged seeds, a second sample 
of seeds (n=1,200) from all six populations was examined (Table 3). The fre- 
quency of normal seeds in a population sample ranged from 1% (population 
#12) to 29% (population #1). 

The next issue addressed was the level of seeds damaged by herbivory vs. 
necrotic seeds (Table 4; n=2,431). Five populations were sampled (all except 
population #8, due to an inadequate amount of seeds). The level of seeds dam- 
aged by herbivory ranged from 10% (population #5) to 38% (population #1). 
The differences between populations in terms of seeds damaged by herbivory 
was not statistically significantly different (ANOVA, F = L9, p = 0.12). For this 
analysis it should be noted that sample size was reduced to 9 plants per popu- 
lation, due to the necessity of having equal sample sizes per population (all 
populations had at least 9 plants with 50 seeds/plant, but not all populations 
had 10 plants with 50 seeds/plant). 

Interspecific comparisons.—Levels of seed herbivory were similar between 
S.shortii population #5 and S. canadensis var. altissima populations #1 +5 (16% 
vs. 18%, respectively) (Table 5). However, the levels of necrosis were significantly 
different—62% for the former, and 19% for the latter (Mann-Whitney Test, U=140, 
p<0.0001). Temporal differences for inviable seed production (combining the 
necrotic and physically damaged groups) between the 2001 and 2003 collec- 
tions (Tables 3 and 5) were not significant for either population #1 (t = 18; 


= 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 741 


TABLE 1.Frequency of VSD (Viable fee Pere comer’ from encheyeley normal seeds sub- 
ject to cold storage as determined by gen a radicle and a complete set of cotyledons 
in a sample of phenotypically normal can 


Pop'n %VSD 


CON NN > 
> 
8 
Ss 


p = 0.081) or population #5 (t = 0.33; p = 0.75). Furthermore, there were no sig- 
nificant differences in inviable seed production between populations #1 and 
#5, as determined by the Mann-Whitney test (U = 130, p= 0.064). In comparisons 
of S. shortii with the local congeners in 2003, the percentage of normal seeds 
was much greater in the more common species (44% and 73% for S. canadensis 
var. altissima for populations #1 and #5, respectively, and 57% for S. nemoralis 
from population #1) (Table 5). Among the damaged seed sets, there was again 
significant S. shortii interpopulation variation for 2003: 36% of the total seed 
sample from population #1 being necrotic but 62% of the population #5 sample 
exhibiting similar necrosis. 

The single Indiana population of Short’s goldenrod exhibited a very high 
level of seed necrosis (74%) in comparison to the Kentucky populations (#1 - 
36%, #5 - 62%). S. gigantea also had a rate of necrosis (41%) higher than that of 
the Kentucky congeners S. canadensis var. altissima (#1 - 21%, #5 - 9%) and S. 
nemoralis (14%). The percentage of normal Indiana seeds for S. shortii (25%) 
was higher than the overall mean among all six Kentucky populations (16%; 
Table 3). In comparing either of the Kentucky S. shortii populations (#1 and 
#5) with the Indiana S. shortii population, there was again no significant dif- 
ference in inviable seed production (Kruskal-Wallis Test statistic = 3.7; p = 0.16). 

There were no differences in inviable seed production between S. shortii 
and S. canadensis occurring syntopically in Kentucky for the comparison in- 
volving population #1 (t-Test, t = 0.14; p = 0.89) but yes for population #5 (t= 5.3; 
p < 0.001) (Table 6). Finally, there were no significant differences in inviable 
seed production between the Indiana population of S. shortii and the syntopic 
congeners S. gigantea and S. nemoralis (Kruskal-Wallis Test statistic = 2.6; p = 
0.2 


The final comparison involved the levels of seed herbivory and necrosis 
among the three populations of S. shortii. The levels of herbivory did signifi- 
cantly differ between Kentucky population #5 and the Indiana population 
(Mann-Whitney Test: U = 85; p = 0.005), but not for population #1 vs. #5 (U = 31; 
p = 0.14) and population #1 vs. the Indiana population (U = 68; p = 0.12). For 


BRIT.ORG/SIDA 22(1 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 743 


Fic. 2. Photomicrographs of the different seed conditions. A = phenotypically normal seeds; B ds subjected to 


7 f \ 
and the general “shrunken” appearance). The bar represents a length of 1 mm. 


necrosis, populations #1 vs. #5 (U = 21; p = 0.025) and population #1 vs. the 
Indiana population (U = 19, p = 0.021) showed statistically significant differ- 
ences. The difference between population #5 and the Indiana population was 
not statistically significant (U = 28; p = 0.087). 


DISCUSSION 


A number of embryo-lethal and mutant developmental alleles have been cata- 
logued (e.g., Meinke 1994), and their existence in the homozygous state at a suf- 
ficient frequency within a population could have extreme consequences for the 
long-term persistence of that population. For that reason, in our tests for seed- 
ling viability, we measured the production of developmentally normal seed- 
lings. In past studies of seed germination (see previously cited Buchele et al. 
and Walck et al.) the primary criterion for determining seed germination was 
emergence of the radicle after two weeks of incubation, as this is the definitive 
physiological indicator for completion of the germination process (Bewley 1997). 
The germination data in these prior studies was then transformed with data 


744 BRIT.ORG/SIDA 22(1) 


Taste 2. Levels of Viable Seed Development (VSD) from a random sample of seeds obtained from 
six populations of Short's goldenrod. All samples consisted of orale normal as well as 
physically damaged and necrotic seeds. ' = data from Smith et a 


Pop’n %VSD Pop’n size! Density! 
] 30% 2,549 0.842 
2 21% 573 0.419 

5 25% 3,488 0.648 
7 14% 1,000 1.91 

8 33% 672 0.877 
12 3% 1,846 0.378 


Taste 3.Proportion of seeds from a random sample that were either phenotypically normal or dam- 
aged (either due to herbivory or necrosis). 


Pop'n % normal % damaged 
] 29% 71% 
2 14% 86% 
5 28% 72% 
7 8% 92% 
8 14% 86% 
12 1% 99% 


Taste 4. Proportion of seed damage due to either external (herbivory) or internal (embryo abor- 
tion?) factors in a sample of damaged seeds from six populations of Short’s goldenrod.. Damaged 
= seeds exhibiting a small hole in the coat, or a broken end; necrotic = seeds with a dark brown or 
black color,and a twisted, shrunken shape. 


Pop’n (sample size) % damaged % necrotic 
| (464) 38% 62% 
2 (487) 30% 64% 
5 (490) 10% 90% 
7 (500) 20% 80% 
12 (490) 12% 88% 


obtained from seeds that had not yet undergone germination, but that contained 
respiring embryos as determined by a histochemical stain (Cottrell 1947). Our 
criteria differ from the criteria previously used to determine seed germination, 
as we posed a different question. 

Our interest in determining the frequency of developmentally normal seed- 
lings produced from each sample of seeds was in obtaining an estimate of com- 
parative reproductive potential. Those populations producing a higher fre- 
quency of developmentally normal seedlings would have a greater potential 
for persistence than a population with-a low level of normal seedlings. Our 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 745 


TABLE zp ECU SINO! He renotypicall normal vs.damaged seeds in two populati f Short's gold- 

ntaining gener species. Nor = phenotypically normal; eget cece exhibiting damage 
die to insect Karbon) Nee = seeds with a darkened and physically distorted evcial appear- 
ance. Populations #1 and 5 are from Kentucky; IN = the Indiana population. The parenthetical nu- 
merals refer to the number of plants sampled. 


Pop’n Taxon Seed condition 

Nor er Nec 
1 S. shortii (10) 54% 10% 36% 
1 S.canadensis var. altissima (5) 44% 35% 21% 
] S.nemoralis (5) 57% 29% 14% 
5 S. shortii (10) 22% 16% 62% 
5) S.canadensis var. altissima (10) 72% 9% 19% 
IN S. shortii (10) 25% 1% 74% 
IN S. gigantea (5) 18% 41% 41% 
IN §.nemoralis (5) 43% 45% 12% 


Taste 6. Summary statistics (mean + 1 SD and range) for the number of inviable (necrotic + dam- 
aged) seeds from a single sample per plant for Kentucky (#1 and #5) and Indiana (IN) 5. shortii and 
Kentucky (#1+#5) S. canadensis populations. 


Pop'n Taxon Mean Range 
] S. shortii 12-263 3-20 
5 S. shortii 15+ 40 8-20 
IN S. shortii 15 241 5-19 
145 S. canadensis LAE 6.0 0-20 


specific question was not one of seed germination rates among and between 
populations, but one of the frequency of normal seedling production. We em- 
ployed incubation conditions that would, based on previous studies (e.g. Walck 
et al. 1997a) optimize seed germination levels, then examined seedlings for nor- 
mal early development. 

During the course of this study, of considerable surprise to us were two dis- 
coveries: the frequencies of seed predation (herbivory) and necrotic (presum- 
ably embryo aborted) seeds. All seven sampled populations experienced marked 
levels of either mechanically damaged seeds or necrotic seeds. The number of 
affected seeds (either damaged or necrotic) in our samples ranged from 70% 
(population #1) to 99% (#12). Of the damaged seed sets, the majority were ne- 
crotic, ranging from 63% (population #1) to 90% (#5). Seed damage in popula- 
tions | and 7 was previously noted “in some infructescences” and attributed to 
larvae of the family Cecidomyiidae (Diptera) (Buchele et al. 1992). There were 
no quantitative data in this earlier report. Another possible candidate is bruchid 
beetle larvae (R. Kingsolver, pers. comm.). 


746 BRIT.ORG/SIDA 22(1) 


There is no previous mention in the literature concerning Solidago regard- 
ing the occurrence of necrotic seeds in mature infructescences. The major threat 
to successtul formation of developmentally normal seedlings appears not to be 
an external agent, e.g.an insect depositing eggs in S. shortii seeds asa food source 
for developing larvae, but rather the frequency of embryo-aborted seeds. There 
could be three possible explanations for the phenomenon of embryo abortion. 
First, due to the small sizes of the populations (Smith et al. 2004) it is possible 
that through stochastic events lethal allele combinations have arisen (Levin 
2000) possibly leading to embryo abortion. Second, there could be pollen com- 
petition among Short’s goldenrod plants and the more common goldenrod spe- 
cies. There is a temporal overlap in anthesis between the three goldenrod spe- 
cies at Blue Licks, and we have observed individual insects visiting 
inflorescences of all three goldenrod species. Likewise plants of the Indiana 
population of Short’s goldenrod could have been pollinated with pollen from 
either S. nemoralis or S. canadensis, leading in both cases to abortive embryo 
formation due to gametic incompatibility (Grant 1971). Third, as Solidago is an 
obligate outcrosser (Buchele et al. 1992) lack of suitable crosspollination would 
lead to unfertilized embryo sacs and arrested development of the resultant seeds 
(Proctor et al. 1996). At present there are no data available to distinguish among 
these three possibilities. 

Itis difficult to assess the impact of reduced seed production on the persis- 
tence of S. shortii, given that three more widespread congener species (S. 
canadensis var. altissima, S. nemoralis and S. gigantea) are likewise subject to 
seed predation. The presence of congener species with a lower level of necrotic 
seeds (and assumedly a pre-predation higher level of normal seeds) does not 
appear to influence the behavior of the seed predators on Short’s goldenrod, as 
there was no statistically significant difference between populations that con- 
tained (#1) or were adjacent to (#5 and the Indiana population) congener spe- 
cies and those without. In the absence of interspecific competition, Walck et al. 
(1999a) observed successful establishment of Short’s goldenrod seedlings in field 
plots over a three-year period. Given the large number of florets in a typical 
Short’s goldenrod inflorescence, there evidently are a sufficient number of vi- 
able seeds generated to provide for some level of sexual reproduction. 


— 


ACKNOWLEDGMENTS 


The able field and laboratory assistance of Ms. Tiffany Carpenter and Ms. Jackie 
Hucul are gratefully appreciated. Thanks are extended to Ms. Deborah White 
of the Kentucky State Nature Preserves Commission and the Kentucky State 
Parks Commission for permission to collect plant materials. Funding was pro- 
vided to PJC through an EKU Faculty Research Award, and from NIH/NCRR/ 
KBRIN Award #1-P20 RR 16481. The review and comments of Dr. Bruce Ford 
on an early version of this manuscript are gratefully acknowledged. 


CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 747 


REFERENCES 


Baskin, J.M.,J.L.Watck, C.C. Baskin, and D.E. Buchete. 2000. Ecology and conservation biology 
of the endangered plant species Solidago shortii (Asteraceae). Native Pl. J. 1:35-41. 

Bewtey, J.D. 1997. Seed germination and dormancy. Pl. Cell 9:1055-1066. 

BucHete, D.E., J.M. Baskin, and C.C. Baskin. 1989. Ecology of the endangered species Solidago 
shortii. |. Geography, populations, and physical habitat. Bull. Torrey Bot. Club 116:344- 


Buchete, D.E.,J.M. Baskin, and C.C. Baskin. 1991.Ecology of the endangered species Solidago 
shortii. Ill. Seed germination ecology. Bull. Torrey Bot.Club 118:288-291 

Buchete, D.E., J.M. Baskin, and C.C. Baskin. 1992. Ecology of the endangered species Solidago 
shortii. lV. Pollination ecology. Bull. Torrey Bot. Club 119:137-141. 

Cortrett, HJ. 1947. Tetrazolium salt as a seed germination indicator. Nature 159:748, 

Evans, M. 1987. Field survey for Short's goldenrod (Solidago shortii). Report for the USFWS 
Region 4, Atlanta, GA. 

Grant, V. 1971. Plant speciation. Columbia Univ., Press, NY. 

Hickox. L.G. and T. Warne. 1998. C-Fern manual. Carolina Biological Supply Co., Burlington, 


Homoya, M.A.and D.B. Asrett. 2005.A natural occurrence of the federally endangered Short's 
goldenrod (Solidago shortii T. & G.) [Asteraceae] in Indiana:lts discovery, habitat, and 
associated flora. Castanea 70:255-262. 

Levin, D.A. 2000. The origin, expansion, and demise of plant species. Oxford Univ. Press, 
New York. 

Meinke, D.W. 1994. Seed development in Arabidopsis thaliana. In Arabidopsis, C.R. Somer- 
ville ,E.M. Meyerowitz, Eds. Cold Spring Harbor Press, NY. 

Smith, B.D., A.J. DenHAM, J.B. Beck, and PJ. Catt. 2004. High resolution GIS mapping and cur- 
rent status of the ten viable populations of Short’s goldenrod (Solidago shortii — 
Asteraceae) in Kentucky. Sida 21:1121-1130. 

Proctor, M., P. Yeo, and A. Lack. 1996. The natural history of pollination. Timber Press, Port- 
land, OR. 

Watck, J.L., JM. Baskin, and C.C. Baskin. 1997a. A comparative study of the seed germination 
biology of a narrow endemic and two geographically-widespread species of Solidago 
(Asteraceae). 1. Germination phenology and effect of cold stratification on germina- 
tion. | Sci. Res. 7:47-58 

Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997b.A comparative study of the seed germination 
aS of anarrow endemic and two g phically-widespread species of Solidago 
(Asteraceae). 2. Germination responses to Ponca seeds in relation to seasona! tem- 
perature cycles. Seed Sci. Res. 7:209-220. 

Watck, J.L., JM. Baskin, and C.C. Baskin. 1997¢.A compalallve study of the seed germination 
Biology of a narrow endemic and two geographically-widespread species of Solidago 
(Asteraceae). 3. Photoecology of germination. Seed Sci. Res. 7:293-301. 

Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997d. A comparative study of the seed germination 


— 


ee) 


748 BRIT.ORG/SIDA 22(1) 


It 


biology of anarrow endemic and two geog ally-widespread species of Solidago 
(Asteraceae). 4. Role of soil moisture in regulating germination. Seed Sci. Res. 7:303- 


Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997e.A comparative study of the seed germination 
biology of a narrow endemic and two g hically-widespread species of Solidago 
(Asteraceae). 5. Effect of dry storage on after-ripening and survivorship. Seed Sci. Res. 
7:311-318. 

Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997f. A comparative study of the seed germination 
biology of a narrow endemic and two geographically-widespread species of Solidago 

Asteraceae). 6. Seed bank. Seed Sci. Res. 8:65—74. 

Watck, J.L.,J.M. Baskin, and C.C. Baskin. 1999a. Effects of competition from introduced plants 
on the establishment, survival, growth and reproduction of the rare plant Solidago 
shortii (Asteraceae). Biol. Conserv. 88:213-219. 

Watck, J.L., JM. Baskin, and C.C. Baskin. 1999b. Relative competitive abilities and growth 
characteristics of a narrowly endemic and a geographically widespread Solidago spe- 
cies (Asteraceae). Amer. J. Bot. 86:820-828. 

Wack, J.L., J.M. Baskin, and C.C. Baskin. 1999c. Ecology of the endangered species Solidago 
shortii.VI. Effects of habitat type, leaf litter, and soil type on seed germination. J. Torrey 
Bot. Soc. 126:117-123. 

Zar, J.H. 1999. Biostatistical analysis, 4" ed. Prentice Hall, Upper Saddle River, NJ. 


— 


NATURALIZATION AND EXTIRPATION OF WATER 
HYACINTH (EICHHORNIA CRASSIPES, PONTEDERIACEAE) 
IN SOUTHWESTERN ARKANSAS, U.S.A. 


Renn Tumlison and Brett Serviss 


Department of Biology 
Henderson State University 
Arkadelphia, Arkansas 71999, U.S.A, 


ABSTRACT 


Southern a pcaae to be at he edge of the eee range for naturalization of the water 
hyacinth, Ei Ims, due to the cold intolerance exhibited by the plant. A 
population discovered eonine in DeGray Lake, a County, Arkansas, was examined from 2002- 


2003 to determine whether the population could survive after the freezes common during winter. 
The population was of particular interest because it had become rooted and remained submerged 
with higher lake levels during the surface freezes of winter. Rosettes that fragmented from the popu- 
lation established new growth during the next growing season, but fluctuating lake levels eventu- 
ally stranded this growth and caused extirpation of the population 


RESUMEN 


Arkansas meridional parece estar en laf lel ial para | lizacion del jacinto 
H 4 ° ] y BES Rae | 

por la planta. 
el Lago DeGray, Condado de Clark, Arkansas, se examino 


de agua, Fichhornia i (M Sol de aan intolerancia 


Una poblacion se descubrié iend 


entre 2002-2003 ae ane si ls poniacon podria sobrevivir después de las heladas comunes 


] 


durante el invierno. La } poe habia llegado a arraigar y quedaba 
d 


ae 4 lr 


sumergida con ie 
Rosetas que se fragmentaron de la poblacion tuvieron crecimiento nuevo en la siguiente estacion de 


de lago cme la la superficie en invierno. 


crecimiento, pero los niveles fluctuantes del lago desfavorecieron finalmente este crecimiento y 
causaron la extincion de la poblacién. 


INTRODUCTION 


Eichhornia crassipes (Martius) Solms (water hyacinth) is one of two natural- 
ized species of Fichhornia found in the United States (Horn 2002; Wunderlin 
1998). Water hyacinth is native to tropical South America, specifically 
Amazonia (Barrett & Forno 1982). Water hyacinth now occurs in over 50 coun- 
tries across five continents (Barrett 1989). It is believed that Eichhornia was in- 
troduced into the United States at a Centennial Exposition at New Orleans in 
1884 (Penfund & Earle 1948). It is one of the most aggressive aquatic weeds in 
the tropics, doubling its population as quickly as every 11-18 days (Penfund & 
Earle 1948). Further, Eichhornia is known to produce physiological changes to 
the aquatic environment (Center & Spencer 1981; Fitzsimmons & Velljos 1986; 
Penfund & Earle 1948; Ultsch 1973). Seed production in temperate regions is 


SIDA 22(1): 749 — 754. 2006 


750 BRIT.ORG/SIDA 22(1) 


usually only about one-half of that in the tropics, primarily because of differ- 
ences in levels of insect visitation (Barrett 1980). 

Fichhornia crassipes is only marginally cold hardy and is completely de- 
stroyed when the rhizome is exposed to temperatures at or below -5°C for 12 or 
more hours (Penfund & Earle 1948). Water hyacinth should not reliably establish 
in areas where these temperature conditions occur annually. In southwestern 
Arkansas, air temperatures reach this level for several days during most winters, 
which likely would limit the yearly regrowth of populations of Eichhornia. 

However, a population of Eichhornia crassipes was discovered in a cove of 
DeGray Lake, Clark County, Arkansas on 28 January 2002. Living rosettes with 
broken stolons were discovered along the north shoreline, and a search to re- 
veal the source of these usually floating plants led to the discovery of the popu- 
lation submerged in the lake waters. We followed the fate of this population, to 
determine whether the submerged condition would allow survival through the 
winter due to protection from freezing. lt was presumed that the typically clear 
water would allow adequate photosynthesis to carry the population until the 
next growing season. 


— 


METHODS 


The source of the original established population was not known. Upon fur- 
ther search, the only locality found supporting plants was the initial discovery 
along the north shore of the cove. Prevailing winds from the south presumably 
pushed floating rosettes into the north shore of the cove and effected establish- 
ment at the site. We assumed that the population was established during the 
previous summer. Numerous old inflorescences were discovered still attached 
to the rosettes of the population. 

We returned to the site on several occasions during 2002 to determine the 
fate of the population. Water temperature (surface and at one meter) and depth 
of the population was measured when possible. 

During 2002, several rosettes collected from the field site were grown in 
the lab and in a small plastic outdoor pond to determine whether the plants 
would flower and produce viable seed. With the onset of winter 2003, a few of 
these rosettes were planted in pots and submerged to determine whether such 
overwintering plants would produce break-away rosettes that would establish 
a new population during spring. 


RESULTS 


The submerged lake population on 28 January 2002 had rosettes of 15-20 cm 
width. The water depth at the location of the deepest specimens (the water in 
the cove was perpetually clear, allowing us to see the plants) was 90 and 107 
cm, but most rosettes were located at a depth of 80-85 cm. The widest portion 
of the population (about | m wide) was located toward the head of the cove, 


751 


where the slope of the substrate was more gradual and the water was shallow, 
but most of the population occurred in a strip about 1/2 m wide by 80 m long 
along the contour of the shoreline. Samples of the rosettes showed old inflores- 
cences. 

Plants toward the head of the cove exhibited morphology more typical of 
floating rosettes, but most other specimens had small floats and an upright 
elongate central or main stem. Temperature at the surface on 28 January was 
12°C, and at one meter was 10°C. 

On 15 February, the population looked as it had previously, but the lake 
had risen about 25 cm so that most plants were at a depth of 105-110 cm. Sedi- 
ments occasionally produced by wave action were deposited as a film over the 
population, although sampled plants still were green. Many plants had become 
separated from the submerged mat and were floating near shore. Temperature 
was 9°C at the surface and 8°C at one meter. 

On 24 April, water depth had increased again, making the rosettes impos- 
sible to see. Several rosettes that had separated and floated now were rooting 
into the sediments along the elevated shoreline, and 10 of these were marked to 
follow their survival. Several green rosettes were trapped among debris at the 
head of the cove, protected from waves created by boaters. Also, eight rosettes 
were found in the last 15 m of the cove along the southern shoreline. Tempera- 
ture was 23°C at the surface and 21°C at one meter. 

By 9 May, water depth had increased again, placing the population at an 
estimated 1.5-2 m depth. The plants rooted into the sediments and marked dur- 
ing the previous trip now were submerged and not relocated. Along the shore- 
line, we counted about 40 rosettes that recently had broken free of the original 
population, many at the head of the cove. One of these was partially buried in 
the wet sandy soil about 1.5 m from water. Most of these stranded rosettes were 
small (3-10 cm width), but had new growth and new roots were evident. Tem- 
perature was 23°C both at the surface and at one meter. 

On 12 June, the highest water level we observed kept the original popula- 
tion, if still alive, submerged beyond visibility. Wind action had placed all float- 
ing rosettes (the 40 observed on 9 May) along the shore near the head of the 
cove. These plants now were larger (8-20 cm width), had new growth, and in- 
cluded clusters with side branches. The specimen partially buried in the sand 
on the previous date was not relocated, and all plants found were floating with 
the exception of a few stranded at the head of the cove by wave action. Tem- 
perature was 31°C at the surface and 29°C at one meter. 

On 18 July, depth of the lake had decreased about 3/4 mand the water was 
very clear, but the original population of Eichhornia had disappeared. Only 18 
rosettes of the 40 were surviving. These were found in 8 clusters all near the 
head of the cove in muddy to sandy areas covered by organic debris and in 
almost constant shade. The roots of these plants had become anchored in sedi- 


752 BRIT.ORG/SIDA 22(1 


= 


ments, and now had rosettes up to 30 cm in diameter and side branches with 
upright leaves with small bladders. These plants were in wet soil located 5-9 m 
from the water of the lake. Temperature was 29.5°C at the surface and 29°C at 
one meter. 

On 9 August, the few remaining exposed plants were drying and dying back, 
with the healthiest three clusters located 5, 5, and 22 m from the lake in the 
sandy soil of the head of the cove. Rains fell before 6 September, which appar- 
ently rejuvenated the plants and permitted new growth of leaves. Additional 
rain fell before a 20 September visit, and had resulted in continued new growth. 

On 7 January 2003, the site was revisited with the expectation that the frost 
of 25 November would have killed the remaining plants that had been isolated 
in exposed locations along the cove. Of the last 18 rosettes, only two were relo- 
cated - dead and submerged under 20 cm of water. 

In the small pond experiment, the rosettes proliferated and filled the pond 
during the summer, and produced numerous inflorescences from 6 August 
through September. However, all surface plants were killed with the first frost. 
The frost killed only exposed portions of 5 otherwise submerged plants. The re- 
maining portions of these 5 plants remained green until early spring, then died. 
No new rosettes were able to break away, surface, and proliferate during the next 
spring. Seeds left in the pond and those used in greenhouse experiments failed to 
germinate within one year, and no germination of seeds has been discovered at 
the field location. It is important to note that the seeds used in our experiments 
were not intentionally scarified to promote laboratory germination. 


DISCUSSION 


A population of Eichhornia crassipes became established in DeGray Lake, at a 
northern latitude presumably marginal for this cold-intolerant plant to over- 
winter outdoors. The original established population survived through periods 
of freezing because the root system anchored the plants, not allowing them to 
float as the level of water in the lake rose during winter. Although the original 
population did not survive the entire winter, rosettes that periodically broke 
free and floated survived if they surfaced after the last freeze. 

The higher level of water during spring deposited these surviving rosettes 
higher up the bank than the original population. Some survived and grew dur- 
ing the next season, but they became exposed and isolated from the lake as water 
levels dropped during summer. Because none of the originally submerged plants 
were observed after 18 July, none was present for future re-establishment. 

From the available data, we believe that the original population was estab- 
lished during the summer of 2001 when the lake level was near its seasonal 
low. Plants grew along the shoreline, held there by the winds, and became an- 
chored by their roots. Any plants that grew across the deeper water of the cove, 
and were not held by the roots as the water level rose, would have died with 


freezing air temperatures. This scenario would have left the observed 80 m long, 
0.5-1 m wide population that followed the contour of the shoreline. This por- 
tion of the original population actually survived the winter by being submerged, 
and re-established and grew the following spring. However, fluctuating lake 
levels led to the extirpation of the population that had established. 

Penfund and Earle (1948) unexpectedly observed that Eichhornia crassipes 
plants died while submerged during the winter. They felt that protection from 
the freezing sur ures would protect the plants, but argued that a lack 
of oxygen might have explained the death of their submerged population. Be- 
cause even clear water filters light, we argue that insufficient light to the sub- 
merged plants might have precluded adequate photosynthesis. Further, water 
saturates with much less oxygen and carbon dioxide than is available in air, which 
likely contributed to the observed death of the rooted population in our study. 

Eichhornia crassipes has a means of survival when a rooted population 
becomes inundated. Penfund and Earle (1948) found that 10 days after submer- 
gence, E. crassipes begins to produce an abscission layer across the rhizome just 
below the lowest living leaves, forms new roots just above the abscission layer, 
and floats to the surface. Those new rosettes could continue the existence of 
the population if surface conditions were within the range of tolerance. 

Seedlings of E. crassipes can survive submergence via the same mechanism 
(Penfund & Earle 1948; Robertson & Thein 1932). Seeds of E. crassipes only ger- 
minate when exposed to air, and thus seeds that germinate on areas where wa- 
ter has receded could become rooted and attached into the soil substrate, and 
subsequently submerged if the water level of the lake rises. We did not observe 
seedling development in our field or lab studies, so we found no evidence that 
re-establishment would occur by those means at our site. 

We believe that this species would be able to survive at our site if the sea- 
sonal high and low water levels were less different, or if the relative timing of 
lake fluctuation and f afforded protection. Thus, submerged plants would 
have a greater chance of surviving longer into the winter and spring, and sepa- 
rating rosettes would not meet high water and be placed too far away from the 
lake to undergo the rapid growth possible during summer. The present condi- 
tions appear to help prevent the long-term establishment of this exotic plant at 
this site. 


ACKNOWLEDGMENTS 


We thank two anonymous reviewers for their comments and suggestions. 


REFERENCES 
Barrett, S. 1989. Waterweed invasions. Sci. Amer. 261:90-97. 
Barrett, S.C. 1980. Sexual reproduction in Eichhornia crassipes (water hyacinth) ||. Seed 
production in natural populations. J. Appl. Ecol. 17:113-124. 


BRIT.ORG/SIDA 22(1) 


Barrett, S.C. and |W. Forno. 1982. Style morph distribution in New World populations of 
Eichhornia crassipes (Mart.) Solms—Laubach (water hyacinth). Aquatic Bot. 13:299-306. 

Center, T.D.and N.R. Spencer. 1981.The phenology and growth of water hyacinth Eichhornia 
crassipes (Mart.) Solms in a eutrophic north-central Florida lake. Aquatic Bot. 10:1-32. 

Fitzsimons, R.E.and R.H. Vattesos. 1986.Growth of water hyacinth [Eichhornia crassipes (Mart.) 
Solms] in the middle Parana River (Argentina). Hydrobiologia. 131:257-260. 

Horn, C.N. 2002. Pontederiaceae.|n: Flora of North America Association, eds. 2002. Flora of 
North America. Volume 26. Oxford University Press, New York, NY. 

Penrunb, W.T. and T.T. Earte. 1948. The biology of the water hyacinth. Ecol. Monogr. 18: 
448-472. 

RoserTsON, H.F.and B.A. THeIn. 1932. The occurrence of water hyacinth (Eichhornia crassipes) 
seedling under natural conditions in Burma. Agric. and Livestock of India. 2:383-390. 

Uttsch, G.R. 1973. The effects of water hyacinths (Fichhornia crassipes) on the microenvi- 
ronment of aquatic environments. Arch. Hydrobiol. 72:460-473. 

WuNDERLN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida. 
Gainesville. 


FUNGI ASSOCIATED WITH BORRICHIA FRUTESCENS 
(ASTERACEAE): INSECT GALLS AND ENDOPHYTES 


Diane Te Strake, Amy Haddock Keagy, and Peter D. Stiling 


Department of Biology 
University of South Florida 
Tampa, Florida 33620-5200, U.S.A. 


ABSTRACT 


This report compares fungi found in galled and non-galled plants of Borrichia frutescens (L.) DC, 
Asteraceae. Fungi were observed in the plant galls of the midge, Asphondylia borrichiae Rossi & 
Strong. A variety o endophytes unely were oe me the apic cal meristems, stems and leaves of 
nels eda nd non-g it Florida. Fifteen percent of midges 


r 
<amin d (n= 60) carried fungal spores of Alternaria sp. 


RESUMEN 
E ticul losh trados en plantas con y sin agallas de Borrichia frutescens 
(L.) DC, Asteraceae Be observaron hongos en las agallas de Asphondylia borrichiae Rossi & Strong. 
Se cultivé una i le los meristemos apicales, tallos y hojas de plantas con y 


sin agallas colectadas en varios lugares costeros de Florida. El quince por ciento de las agallas 
examinadas (n = 60) tenian esporas de Alternaria sp. 


INTRODUCTION 


Borrichia frutescens (L.) DC. or bushy seaside oxeye is a perennial herbaceous 
Asteraceae often found bordering the salt marsh communities of the Gulf of 
Mexico and Atlantic Ocean. Insect galls on this host species are initiated in the 
apical meristem by a midge, Asphondylia borrichiae Rossi & Strong (1990). 
These galls are also impacted by four specific parasitoid wasps (Galeopsomyia 
haemon Walker, Rileya cecidomyiae Ashmead, Tenuipetiolus teredon Walker, 
and Torymus umbilicatus Gahan). Individuals from these parasites lay their eggs 
inside the gall with A. borrichiae. The parasitoids devour the developing midges 
and significantly affect population levels of A. borrichiae (Rossi et al. 1992). 
Several biological studies on this plant midge system elaborate on the life his- 
tory and ecological details including parasitism rates (Rossi & Strong 1990; 
Stiling et al. 1992; Stiling 1994; Rossi & Stiling 1998; Rossi et al. 1999). Asin many 
insect galls, the tissue surrounding the developing larvae eventually becomes 
full of chambers layered with fungal growth purportedly providing nutrition 
for the developing larvae (Gagne 1989). Thus, knowledge of the fungi associ- 
ated with both the galled and non-galled plants is needed to gain insight into 
this host-parasite relationship. 

Itis unclear how fungi are introduced into the galls in the B.frutescens sys- 
tem. Many plants are known to harbor microbial endophytes which predomi- 


SIDA 22(1): 755 — 763. 2006 


756 BRIT.ORG/SIDA 22(1) 


nantly are fungi. These organisms live internally in plant tissue. Stone et al. 
(2000) suggest that these endophytes secondarily invade insect galls. Batra and 
Lichwardt (1963) indicate that “gall fungi are airborne, grown in a variety of 
substrata and are not species specific.” Other investigators of insect galls sus- 
pect that spores of fungi are accidentally collected from leaves and leaf litter 
by the insects (Borkent & Bissett 1965). Haridass (1987) suggests that fecal con- 
tamination is another possible means of spore dispersal whereby fungi are in- 
troduced into the plant tissue. Gagne (1989) reports that midges feed by suck- 
ing on the hyphae and states, “fungi are evidently obligatory in all galls..and 
are food of those gall midges.” Batra and Lichwardt (1963) alternatively indi- 
cate that it is difficult to determine whether midges actually feed on the hy- 
phae or the plant tissue degraded by the fungi. 

In this study we examined a number of midge galls of Asphondylia 
borrichiae from Borrichia frutescens from several Florida sites for the presence 
or absence of fungi. In addition plant apices, stems, and leaf samples of galled 
and non-galled plants were cultured for fungi. Finally, a small population of 
midges was surveyed for associated fungal spores. 

MATERIALS AND METHODS 
Sites for Collections.—Borrichia frutescens galls, leaf and stem samples were 
collected from the following locations from 1993-1995 in Florida: 

Site #1-Merritt Island National Wildlife Refuge, Titusville (Brevard 
County). 

Site #2-Little Jetties Park, Mayport (Duval County). 

Site #3-Ft. DeSoto State Park, St. Petersburg (Pinellas County) 

Site #4-Upper Tampa Bay Park (Hillsborough County). 

Site #5-Delnor-Wiggins State Park, Naples (Collier County). 

Culture Protocol.—In May, 1995, five samples each of apex, stem, and leaf tissue 
were collected from galled and non-galled plants from Sites 1, 2, and 3. All col- 
lected plant samples were placed in individual, sterile plastic bags, refrigerated, 
and returned to the laboratory within three days. Excised tissue from apices, 
stem and leaf tissue of both galled and non-galled plants tissues were rinsed in 
running water for 5 min, then each tissue type was placed in separate flasks 
and rinsed with 95% ethanol for 2 min. This was followed by a 30 min shake 
rinse in 20% bleach solution. All tissues were held in flasks of sterile distilled 
water (1- 2h) while tissues were prepared for incubation. Three slices from each 
tissue [rom each site were submerged in Potato Dextrose Agar (PDA) and V-8 
Juice Agar (Difco, Detroit, MI). These cultures were placed in a dark incubator 
at 32°C for seven days. Isolates from successful cultures were placed on PDA 
slants and held at ambient temperature until sporulation after which identifi- 
cations were made. 


a 


TE STRAKE ET AL 757 


In Situ Fungal Studies.—In September, 1995, ten stems were collected ran- 
domly from galled and non-galled plants of B. frutescens from four sites (# 1- 
4). Each stem was cut into 1.5 cm segments. Using the protocol established by 
Hignight et al. (1993) stems were cleared and then all segments were micro- 
scopically examined for the presence of endophytes. 

Direct Observations of Insects.— Whole insects of Asphondylia borrichiae 
were collected as they emerged from galls of Borrichia frutescens, (n = 60), Iva 
imbricata Walter (n = 20), and Iva frutescens L. (n =5) and preserved in vials of 
95% ethanol. All insect samples were examined for the presence or absence of 
fungal spores on the surface using light microscopy. 


RESULTS AND DISCUSSION 


Previous dissection of the Borrichia frutescens galls revealed larval chambers 
(pers. obs.). Within these white mycelium could be observed and this fungal 
growth was arrested until the larvae became fully developed. Then the myce- 
lial mat became denser, darker, and quite distinct (Fig. 1). 

Midge galls observed in the Asteraceae, and fungi isolated from those galls 
are listed, in Table L. In addition, Farr et al.(1989) note the occurrence of Aecidium 
borrichiae, Puccinia mirifica, and Cercosporidium sp. on stems and leaves of B. 
frutescens. Although many plants have fungal endophytes (Clay et al. 1985; Clay 
1990; Carroll 1988), prior to this they were not known from B. frutescens. 

Preliminary studies (1993-1994) including direct observations and excis- 
ing and culturing of a range of ages of gall tissue from B. frutescens from all five 
sites consistently yielded a large variety of fungi including two types of sterile 
mycelium (data not shown). Similar observations have been made in other stud- 
ies of midge galls. Bissett and Borkent (1988) ogee that some midges from the 
hondylidii late galls witha variety of mitosporic fungi 
and use them as a food source. Stone et al. (2000) observed that galls on Dou- 
glas-fir (Pseudotsuga menziesii) support heavy fungal growth which, he sug- 
gests, may be plant pathogens in addition to providing insect nutrition. Wilson 
(1995) proposes that fungi in galls may be saprobes or inquilines (organisms 
inhabiting insect galls not parasitizing the gall maker but otherwise utilizing 
the gall tissue for food). 

To determine if endophytic fungi could be observed microscopically and 
using the method of Hignight et al. (1993), a preliminary study was conducted 
in May 1995 using five stems each from galled, non-galled, flowering non-galled 
and flowering galled plants of B. frutescens collected from Site #3, Ft. Desoto. 
This mini-study revealed that endophytic fungi were present in 100% of the 
samples from plants with galls appearing similar to the example seen in Figure 
2. The remaining non-galled stems were less heavily colonized with fungi. This 
qualitative observation led to additional studies to quantify endophytic fungi 


Lasiopterini and A 


758 BRIT.ORG/SIDA 22(1) 


involved in the stems of galled and non-galled plants. We observed a consistent 
occurrence of endophytic fungi in all sampled stem segments from four diverse 
Florida locations (Sites 1-4). In all non-galled plants these occurrences range 
from 20 to 64% and in galled plants 30 to 64%. This suggests little influence of 
the role of the midge. But one might argue that the fungal endophytes in galls 
grow out, invading other plant tissue. On the other hand, depending on the time- 
frame, endophytic fungi may grow into the gall or exist in the pre-gall tissue 
and contribute to the mycoflora of the aging gall. Fither way, fungal endophytes 
are extant in B. frutescens. 

In the apices, stems, and leaves of both galled and non-galled plants from 
Sites l, 2, and 3 several fungal taxa occur (Table 2). Included among the isolates 
are several types of sterile mycelium. These were found among all tissue types. 
Alternaria sp. was also found among all tissue types (Table 2). Acremonium strictum, 


TE STRAKE ET AL., FUNGI 


WITH BORRICHIA FRUTESCENS 


Taste 1.Fungi reported in gall midge associations in Asteraceae. 


Gall midge 


Host 


Fungi 


Reference 


Asteromyia carbonifera 


Solidago canadensis 
S. mollis 


Sclerotium asteris 
Macrophoma sp. 


Weis 1982; Batra 1964 
Borkent & Bissett 1985 


A.carbonifera Aster s Rhytisma asterts Batra 1964 
A.carbonifera S. sempervirens R. bifrons Batra 1964 
A.carbonifera S. lanceolata R. solidaginis Batra 1964 
A. ratibidae Ratibida columnifera  Chaetomium globosum Batra 1963 
A. ratibidae R. columnifera Aureobasidium Batra 1963 
pullulans 
A. ratibidae R. columnifera Plectosporium 
tabacinum 
(as Cephalosporium Batra 1963 
ciferrii 
A. carbonifera 5.graminifolia Alternaria sp. Batra 1963 
Bucculatrix simulans Helianthus sp. Trichoderma viride Batra 1963 
B. simulans lianthus sp Aspergillus sp. Batra 1963 
B. simulans Helianthus sp Trichothecium roseum Batra 1963 
B. simulans Helianthus sp Penicillium spp Batra 1963 
B. simulans Helianthus sp Stemphylium sp Batra, 1963 
B. simulans Helianthus sp. Botrytis cinerea Batra 1963 


Bipolarissp.,and Verticillium | ii displayed more irregular distribution in the 
plant tissues from the sites sampled. Other species of these genera have been 
reported in the endophytic fungal literature (Bacon & White 2000). 

The mycangia (small special pockets adjoining the abdomen) of midges in 
the wild are known to be filled with fungal spores that are thought to be in- 
serted with the eggs at oviposition. In the newly emerged midges examined in 
this study, no spores were seen in the mycangia. Spores of Alternaria sp. were 
seen on the surfaces of 15% of the midges emerging from Borrichia frutescens 
galls (n = 60); 10% of the midges from Iva imbricata (n=20), and 40% of these 
midges from I. frutescens(n=5). Limited observations of several epidermal peels 
from plants of B.frutescens revealed that Alternaria sp. and Fusarium sp. were 
abundant on the plant surface. While it may be possible for the midges to carry 
fungal spores at oviposition, it seems that the diversity of fungi seen in these 
midge galls and host plant tissue are sourced by other means, possibly includ- 
ing some of their parasitoids. 

We observed that fungal endophytes appear to become denser in the plant 
tissues between May (seen in the preliminary study) and September revealing 
increasing hyphae and pigmentation (Fig. 2). The aging galls in this study were 
seen to include fungal contents that became denser and darker with pigments 
(Fig. 1). Alternariasp.and Bipolarissp., both darkly pigmented, were consistently 


760 BRIT.ORG/SIDA 22(1) 


Taste 2.Percent occurrence of fungi isolated from apical, stem and leaf tissues isolated from Borrichia 
frutescens: galled and non-galled plants. 


Merritt Island Mayport Ft Desoto 
gall non-gall gall —_non-gall gall non-gall 


Sterile mycelia 


Verticillium lecanii 

(A. Zimmerm.) Viegas 
Apex - = - - = 
Stem = a 67 - a _ 
Leaf - 20 - - 20 ~ 
Bipolaris sp. 
Apex - 7 - = 7 
Stem 6.7 ~ = - = - 


Acremonium strictum W.Gams 
pex - - - 
Stem 13.3 20. == - 
Leaf - 7 - - - - 
Humicola grisea Traaen 
Apex - = - — 
Stem - - 6.7 ~ 
eaf — - - ~ 


Monocillium indicum 
S.B. Saksena 


Apex - 
Stem = = = = 6.7 & 
Leaf - - - - = . 


Penicillium sp. 
x 


Fusarium sp. 

pex - - - - 
Stem 6.7 7 ~ = - - 
Leaf - - - - - ~ 


TE STRAKE ET AL.. FUNGI ASSOCIATED WITH BORRICHIA FRUTESCENS 161 


isolated from galled and non-galled plant tissue. Shaw (1992) reviewed several 
studies indicating that fungivores consistently favored darkly pigmented litter 
fungi. Interestingly, it seems that Alternaria sp. and Bipolaris sp. could play a 
nutritional role in this gall system. 

In this study we establish the presence of entophytic fungi in insect galls. 
Endophytic fungi were cultured from tissues of galled and non-galled plants 
of Borrichia frutescens. Both of these plant types carry several fungal taxa. These 
observations are common toall sites, but sample sizes are too limited to discern 
major differences. Fungal spores were directly observed on 15% of examined 
midges. This suggests that the midges may play a role in fungal dispersal to gall 
tissue as proposed by Batra and Lichtwardt (1963). 


ACKNOWLEDGMENTS 
The authors greatly appreciate the comments and suggestions of Joe Hennen, 
Erica Cline, and one anonymous reviewer. 
REFERENCES 
Bacon, C.W. and J.F. Waite, Jr. (eds.) 2000. Microbial endophytes. M. Dekker, Inc., New York. 


762 BRIT.ORG/SIDA 22(1) 


Batra, L.R. 1964. Insect-fungus blister galls of Solidago and Aster. J. Kansas Entomol. Soc. 
37:227-234. 

Barra, L.R.and R.W. LichTwaror. 1963. Association of fungi with some insect galls. J. Kansas 
Entomol. Soc. 36:262-278. 

Bissett, J. and A. Borkenr. 1988. Ambrosia galls: The significance of fungal nutrition in the 
evolution of the Cecidomyiidae (Diptera). In: K. Pirozynski and D. Hawksworth, eds. Co- 
evolution of fungi with plants and animals. Academic Press, New York. 

Borkent, A. and J. Bisset. 1985. Gall midges (Diptera: Cecidomyiidae) are vectors for their 
fungal symbionts. Symbiosis 1:185-194. 

Carroll, G.C. 1988. Fungal endophytes in stems and leaves from latent pathogen to mu- 
tualistic symbiont. Ecology 69:2-9. 

Cay, K. 1990. Fungal endophytes of grasses. Ann. Rev. Ecol. Syst. 21:275-297, 

Ctay, K.,T. Harpy, and A.M. HamMmonp, Je. 1985.Fungal endophytes of grasses and their effects 
on an insect herbivore. Oecologia 66:1-5. 

Farr, F., G. Bits, G. CHameris, and A.Y. Rossman. 1989. Fungi on plants and plant products in 
the United States. American Phytopathological Press, St. Paul. 

Gacne, R.J.1989. The plant-feeding gall midges of North America. Comstock 
PublisherAssociates. Ithaca, NY. 

Haripass, E.T. 1987, Midge-fungus interactions in a curcubit stem gall. Phytophaga 1: 
57-74. 

HicnicHT, K., G.A. Muiensure, and A.J.P. van Wuk. 1993.A clearing technique for detecting the 
fungal endophyte Acremonium sp.in grasses. Biotechnic & Histochemistry 68:87-90. 

Rossi, A. M. and D.R. Strona. 1990. A new species of Asphondylia (Diptera:Cecidomyiidae) 
on Borrichia (Asteraceae) from Florida. Proc. Entomol. Soc. Washington 92:732-735. 

Rossi, A.M., P.D. Stitinc, D.R. STRONG, and D.M. JoHNson. 1992. Does gall diameter affect the 
parasitism rate of Asphondylia borrichiae (Diptera: Cecidomyiidae)? Ecol. Entomol. 
17:149-154. 

Rossi, A.M. and PD. StiLinc. 1998. The interactions of plant clone and abiotic factors ona 
gall-making midge. Oecologia 116:170-176. 

Rossi, A.M., P.D. StiLinc, M.V.Cartett, and T.|. BowpisH. 1999. Evidence for host-associated races 
in a gall-forming midge: tradeoffs in potential fecundity. Ecol. Entomol. 24:95-102. 
SHaw, P.1992. Fungi, fungivores, and fungal food webs.|n:G.C. Carroll and D.T.Wicklow, eds. 
The fungal community: its organization and role in the ecosystem. M. Dekker, Inc., New 

York. Pp. 295-310. 

STILING, P.D., A.M. Rossi, D.R. StRoNG, and D.M. JoHNSON. 1992. Life history and parasites of 
Asphonadylia borrichiae (Diptera: Cecidomyiidae), a gall maker on Borrichia frutescens. 
Florida Entomol, 75:130-137. 

STiLinG, PD. 1994, Coastal insect herbivore populations are strongly influenced by envi- 
ronment variation. Ecol. Entomol. 19:39-44. 

Stone, J.K., C.W. Bacon, and J.F. Wuite, Jr. 2000. An overview of endophytic microbes: 
endophytism defined. In: Bacon, C.W. and J.F. White, Jr, eds. Microbial endophytes. M. 
Dekker, Inc., New York. 


= 


TESTRAKE ET AL., 763 


Weis, A.E. 1982. Use of a symbiotic fungus by the gall maker Asteromyia carbonfera to 
inhibit attack by the parasitoid Torymus capite. Ecology 63:1602-1605. 

Witson, D. 1995. Fungal endophytes which invade insect galls: insect pathogens, benign 
saprophytes, or fungal inquilines? Oecologia 103:255-260. 


BRIT.ORG/SIDA 22(1) 


Book REVIEW 
Eucene N. Koztorr. 2005. Plants of Western Oregon, Washington & British Co- 
lumbia. (ISBN 0-88192-724-4, hbk.). Timber Press Inc, 133 S.W. Second 
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: 
www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327- 
5680, 503-227-3070 fax.). $65.00, 608 pp., 712 color photos, 354 b/w line 
drawings, 7 3/8" x 10 3/8" 


“This new book, using a relatively small voca 


a.) 


eae y of scientific terms, is designed for the identifica- 
the crest of the Cascades, from southern Oregon north 


tion of vascular plants ... that grow west ¢ 
through southern British Columbia. It is ‘ very useful on the higher portions of the slopes east of 
the crest of the Cascades.” More than 2500 species are included—354 line drawings (interspersed 
through the text, mostly by Jeanne Janish and John Rumely from Vascular Plants of the Pacific North- 
west) and 709 excellent color photos (arranged together in the middle of the book). A brief discus- 
sion is provided for each family, followed by a key to genera, then by keys to the species of each genus. 

shabeti- 


> U 


| morphological and distributional information is conveyed in the keys. All taxa are a 
cally arranged, without authorities and publication details; common names are provided but scien- 
tific synonymy is minimal. T dated from predeces- 
sors, but concepts of families and genera remain conservative. Fore 


ne species coverage and nomenclature is much up 
cample in the sterace 


—~ 


cae groups 


of similar species are keyed in conglomerates (Bal hiza-Wyethia complex, Aste 
Machaeranthera complex, Ericameria-Pyrrocoma- Columbiadori id-Hazardia complex, ies. 
li \ l | lex). The user friendly keys, the 


Euthamia complex, Madia-Ani SOCUFPUus Hemizonella 
heart and soul of this volume, appear to be effective and Kozloft’s Weak undoubtedly will quickly 


find it nto wide use.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort 


Worth, ee ee U.S.A 


SIDA 22(1): 764. 2006 


TYPE LOCALITIES OF VASCULAR PLANTS FIRST 
DESCRIBED FROM OHIO: SUPPLEMENT! 


James S. Pringle 


R yal B tanical Gardens 
PO. Box 399. Hamilton 
Ontario, CANADA L8N 3H8 
Jpringle@rbg.ca 


In 2003 Ronald L. Stuckey and I published a list of Ohio localities at which the 
type specimens of the names of vascular plants had been obtained, including 
the plant names based on specimens from each locality. Omissions have be- 
come apparent, hence this supplement. As in the original list, double asterisks 
denote names that are currently accepted for the respective taxa. Otherwise, 
the currently accepted names are given in brackets. Minor variants not other- 
wise accounted for are now generally not recognized taxonomically, but remain 
included within the same species. Further explanation of the format appears in 
our earlier paper. 


OHIO 

Carex xanthocarpa E.PBicknell, Bull. Torrey Bot. Club 23:22. 1896, not Degl. 1807: 
C. vulpinoidea var. xanthocarpa Kuk., Pflanzenr. (Engler) 4(20, heft 38):148. 1909, 
based on C. xanthocarpa E.PBicknell; C. bicknellii E.G.Camus, Notul. Syst. (Paris) 
1:239. 1910 and C. brachyglossa Mack., Bull. Torrey Bot. Club 50:355. 1923, both 
published in substitution for C. xanthocarpa E.PBicknell [Carex annectens 
(E.PBicknell) E.PBicknell] 

Laciniaria shortii Alexander ex Small, Man. S.E. Fl. 1335.1933|Liatris squarrulosa 
Michx]]. 

Vernonia maxima Small, Bull. Torrey Bot. Club 27:280. 1900 [Vernonia gigantea 
(Walt.) Trel. var. gigantea. 


Maumee RIVER 

Baptisia leucantha Torr. & A. Gray, Fl. N. Amer. 1:385. 1840 [Baptisia alba var. 
macrophylla (Larisey) Isely]. 

Miami River, Miami Country 

Geum franckii [sic] Steud., Nomencl. Bot., ed. 2, 1:683. 1840, not Sm. 1810: pub- 
lished in substitution for Geum parviflorum Hochst. & Steud., that name not 
validly published [Geum vernum (Raf.) Torr. & A. Gray], 


R 


‘Contribution No. 128 from the Royal Botanical lens, Hamilton, Ontario, Canada, 


SIDA 22(1): 765 - 767. 2006 


766 BRIT.ORG/SIDA 22(1) 


FRANKLIN COUNTY 

Columbus. Trillium erectum var. declinatum A. Gray, Man., ed 5, 523. 1867, T. 
declinatum (A. Gray) Gleason, Bull. Torrey Bot. Club 33:389. 1906, not Raf. 1840, 
and T. gleasonii Fernald, Rhodora 34:21. 1932, published in substitution for T. 
declinatum (A. Gray) Gleason [Trillium flexipes Raf]. 

HAMILTON COUNTY 

Cincinnati. Aira controversa Steud, Syn. Fl. Glumac. 1:224. 1854 [Sphenopholis 
intermedia (Rydb.) Rydbl]. 

Crataegus coccinea [var] e? [sic] mollis Torr. & A. Gray, Fl. N. Amer. 1:465. 1840 
[Crataegus coccinea L., not Crataegus mollis Scheele]. 

Lippia nodiflora var. acutifolia Kuntze, Revis. Gen. Pl. 2:508.1891[Phyla nodiflora 
(L.) Greene; often retained in Lippial. 

Stachys cincinnatensis Kuntze, Revis. Gen. Pl. 2:521. 1891 [Stachys tenuifolia 
Willd! 

Volvulus sepium var. biangulo-sagittatus [sic] Kuntze, Revis. Gen. Pl. 2:447. 1891 
[Calystegia sepium (L.) R.Br. subsp. angulata Brummitt]. 

Cincinnati, Lawrence Bay. Sida spinosa var. flava Kuntze, Revis. Gen. Pl. 1:73. 
1891. 

Lawrenceburg. Iriha [sic] autumnalis var. caespitosa Kuntze, Rev. 2:752. 1891 
[Fimbristylis autumnalis (L.) Roem. & Schult]. 


HockinG County 
Cedar Falls. **Vittaria appalachiana Farrar & Mickel, Amer. Fern J. 81:69. 1991. 


LoraAIN COUNTY 
North Amherst. Geranium carolinianum var. confertiflorum Fernald, Rhodora 
37:300. 1935 [Geranium carolinianum L. var. carolinianuml. 


Lucas County 

Alexis [interpreted as the railroad junction in present-day Toledo rather than 
the community in Monroe County]. Cardamine douglasii forma albidula Farw,, 
Amer. Midl. Nat. 9:261. 1925. 

Toledo, Maumee River bank, Walbridge Park. Malus coronaria var. dasycalyx 
Rehd., J. Arnold Arbor. 2:52. 1920. 


Meics County 

Slope above Leading Creek, Co. Rt. 10, 0.25 mi SW of Twp. Rt. 27, N of Dexter, 
Sect. 6, Salem Twp. Polypodium x incognitum Cusick, Amer. Fern J.92:241. 2002 
[= Polypodium appalachianum Hauller & Windham ~ P.virginianum Ll. 
RICHLAND COUNTY 

Mansfield. Convolvulus sepium var. communis R.M. Tryon, Rhodora 41:419. 1939 
[Calystegia sepium (L.) R.Br. subsp. sepium. 


PRINGLE OHIO PLANTS 767 


Scioto COUNTY 
Friendship, Shawnee State Forest, Camp Gordon C.C.C. Disporum schaffneri 
Moldenke, Phytologia 1:167. 1935 [Prosartes maculata (Buckley) A. Gray]. 


In addition, the names *Oxalis grandis Small, based on O. recurva Elliott sensu 
Trelease, not as originally applied by Elliott, and **Sanicula trifoliata 
E.PBicknell were originally based on syntypes that included specimens respec- 
tively from “Ohio” and “Franklin County, Ohio.” Elymus canadensis var. inter- 
media Vasey ex S.Watson @ J.M.Coult. was originally described with no speci- 
mens cited but with the range said to include Ohio [now included in E. virginicus 
L. var. virginicus]. For all of these mames later authors have designated speci- 
mens from other states as the lectotypes. 


REFERENCE 


Stuckey, R.L. and J.S. Princte. 2003. Type localities of vascular plants first described from 
Ohio. Sida 20:1677-1692. 


768 BRIT.ORG/SIDA 22(1) 


Book NOTICE 


Prrer Gregory. 2000. Plant Roots: Growth Activity and Interaction With Soils. 
(ISBN 1-4051-1906-3; 978-1-4051-1906-1, hbk.). Blackwell Publishing Pro- 
fessional, 2121 State Street, Ames, IA 50014-8300, U.S.A. (Orders: 
www black wellplantsci.com, 800-862-6657, 515-292-3348 fax). $199.99 hbk., 
328 pp., illustrations, 7" x 10' 

As suggested by the cover notes, “this book will be a vital tool for plant, crop, soil, and agricultural 

I scientists, ecologists, and hydrologists.” It’s expensive, 


scientists, plant physiologists 
but perhaps “essential” as well as “vital.” There are nine chapters, densely packed with technical in- 
formation but easily readable, as follow. 

. Plants, Roots and the Soil (evolution, roots and shoots, roots and soi 
Architecture of Root Systems (structure, extension ae branching, root tip, system 


. Roots and th 

ees 

socal men a Growth of Root Systems (measurement of root systems, system development, 
ng) 


size, n, longevity and turnover, modelli 


4. The Functioning Root System (anchorage, water ae nutrient uptake). 
5. Roots and the Physico-chemical Environment (temperature, tropistic responses, soil pores anc 


mechanical properties, atmospheric CO2). 
Environment (soil organisms, symbioses, pathogens and parasites, root 


. Roots and the Biologica 
herbivory 

7. The Rhizosphere (rhizodeposition, chemical changes affecting nutrient acquisition, physical 
changes in the rhizosphere) 

8. Genetic Control of Root System Properties (genotypic differences, genetics, breeding). 

9. Root Systems as Management Tools (optimal root systems, intercropping and agroforestry, crop 
rotations, phytoremediation). 

—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A 


SIDA 22(1): 768. 2006 


NOTES ON SOUTHWESTERN MORACEAE 


Alan T.Whittemore 
1/S National Arh retum 
1 New York Ave NE 
ie. DC 20002-1958 U.S.A. 


ABSTRACT 
Specimens forming the basis of reports of Ficus palmata (F pseudo-carica) in North America are 
misidentified and actually belong to F. carica, so F palmata must be excluded from the adventive 
‘lora of North America. Morus nigra is present as a rare waif in ee nia and Nevada. It is best 
distinguished from other Morus species in the area by its deeply cordate leaves and densely pubes- 
cent styles 3-5 mm long 


RESUMEN 


Los especimenes en que se basan las citas de Ficus palmata (E pseudo-carica) en Norte América estan 
mal identificados y pertenecen a F carica, por lo que E palmata debe excluirse de la flora adventicia 
de Norte América. Morus nigra esta presente como una planta rara abandonada en California y Ne- 
vada. Se diferencia de otras especies de Morus en el area por sus hojas profundamente cordadas y 
estilos densamente pubescentes de 3-5 mm de longitud. 


Moraceae are primarily trees and shrubs of tropical and subtropical areas, with 
few herbaceous species and few taxa found in areas with cooler climates. The 
family is poorly represented in the flora of the southwestern United States: re- 
cent floras list only two native species, Morus microphylla Buckl. (which oc- 
curs from central Texas west to Arizona and across northern Mexico) and M. 
rubra L. (an eastern species that crosses the Great Plains in Texas), and three 
introduced species, M. alba L., Ficus carica L., and Maclura pomifera (Raf.) 
Schneid., from the western half of the country (McClintock 1993; Wunderlin 
1997). Two other exotic species not mentioned in these floras, Ficus palmata 
Forrsk. and Morus nigra L., have been recorded as escapes in California and 
Nevada. In the course of revising McClintock’s treatment for the second edition 
of the Jepson manual of the higher plants of California, specimens vouchering 
these reports were examined in order to clarify the status of these species as 
possible escapes in North America. 


California reports of Ficus palmata 

Ficus palmata is a small tree or shrub native from Iran east to India and Nepal, 
and south to Ethiopia and Somalia. It is closely related to the commercial fig, 
Ficus carica L., a native of the Near East that is now naturalized at scattered 
sites across much of the southern United States (Wunderlin 1997), but F palmata 
differs in having smaller, often unlobed, leaves and smaller fruit (Browicz 1982: 
Friis 1993; Ghafoor 1985). Ficus palmata (=F. pseudo-carica Miq.) has been re- 


SIDA 22(1): 769 - 775. 2006 


770 BRIT.ORG/SIDA 22(1) 


ported from southern California, first by Munz (1974). Munz used the newer 
synonym E pseudo-carica Mig. and described it as similar to F carica, but with 
leaves more deeply divided, a description that is at odds with descriptions and 
keys in recent floras of areas where F palmata is native (Browicz 1982; Friis 
1993; Ghafoor 1985). Munz listed the species as “Reported as occasional escape 
in Santa Barbara region,” the phrasing suggesting that Munz picked up his iden- 
tification from another source. Smith (1976, 1998) mentioned two collections 
by Henry M. Pollard from “west fork of Cold Springs [sic] Canyon, Montecito.” 
The species was not treated for California in the most recent state flora (Mc- 
Clintock 1993) or the Flora of North America (Wunderlin 1997), but it was re- 
ported again (this time under the older name F palmata) by Hrusa et al. 2002), 
who cited a single collection from Santa Barbara County, California, made by 
Henry Pollard in 1958. 

In view of the irregular treatment of this species (reported by several local 
floras, but not the most recent state and national floras) and the mismatch be- 
tween the keys and descriptions of F palmata in the North American and Old 
World literature, it seemed best to examine the specimens that the California 
reports are based on and compare them with specimens collected from the na- 
tive range of E palmata in the Old World, as well as the full range of variation 
in cultivated F carica. 


Variation in Ficus palmata and FE. carica 
Thirty-one specimens from three herbaria (MO, NA, and US), representing Fi- 
cus palmata from throughout its native range, were examined and coded for 
morphological characters that potentially distinguish the species from F carica. 
The results are shown in Table 1, along with characters of cultivated F. carica, 
taken from 70 sheets of cultivated figs, and characters of the vouchers for the 
California oS of F. palmata (see below). Characters taken from herbarium 
s agree well with descriptions and illustrations of the two species from 
Este sources (Browicz 1982; Condit 1947; Friis 1993; Ghafoor 1985). 


Representative specimens examined: ses aaa INDIA: Botanical Garden, University of Delhi, 
Rodin 8049 (US). Setabani, Hem Raj s.n. 28 Feb 1931 (MO). Koraput district, Jaypore Estate, Orissa, 
3000-3500 ft, H.E Mooney 3893 (MO). ee a W. Koelz 20496 (NA). Almora, United Prov- 
inces, 6000 ft, W. Koelz 19996 (NA). PAKISTAN: Peshawar, W. Koelz 8206 (NA); 3 mi from Abbotabad 
on road to Havelion, M. Qaiser & A. Ghafoor 1999 (NA), Karakar, M. Shah & Dilawar 1001 (MO). NE- 
PAL: Wada, 6600 ft, Mr Hoh and Rajbhandari 1157 (US). SAUDI ARABIA: Al Karn, Dwyer & El Sheikh 
13676 (MO). YEMEN: Attara, 1900 m, Deflers 402 (US). ERITREA. Amasen: Ad Rassi, eis 4925 (US). 
ETHIOPIA. Kaffa Prov.: 7 km E of oe Burger 1158, Meyer 7820 (US). Harar Proy.: 22 km SE of 
Harar, ca. 1350 m, Burger Ae 1622 (US). 

Ficus carica (all cultivated, a f depos at ay CHILE. Juan Fernandez Islands: Mas Afuera, 
EG. Meyer 9461 IRAN. Tehran, W. Koelz 16085. U.S.A. Arkansas. Drew Co.: Monticello, E. Sundell 91LL 
California. Alameda Co.: es 23041. Butte Co.: Chico, WF Wight 1279. Riverside Co.: Citrus 
Experiment Station, Riverside, LJ. Condit 1. Santa Barbara Co.: Franceschi Nursery, Santa Barbara, 
W.E Wight 4879. Sonoma Co.: Sonoma, H. Grinstead s.n. 4 Jun 1978. Florida. Alachua Co.: Gainesville, 


WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 771 


EG. Meyer & PM. Mazzeo 14365. Georgia. Morgan Co.: Madison, FG. Meyer & PM. Mazzeo 20440. 
Illinois. Cook Co.: Morton Arboretum, Lisle, N. Gavlak 4098V90. South Carolina: Columbia, J.B. Nel- 
son 19580. ZIMBABWE: Salisbury, H. M. Bigel 4444. 
California material referred to Ficus palmata 
The collections cited as Ficus palmata and F. pseudo-carica by Smith (1976) and 
Hrusa et al. (2002) are part of a series of four collections made by Henry Pollard 
(Pollard s n. 20 November 1955, 29 September 1956, 23 December 1958, and 28 
November 1959 [all CAS]). All bear the same locality data (bed of west fork of 
Cold Spring Canyon, Santa Barbara Co.) and evidently came from the same 
colony (perhaps the same plant), described by Pollard on the herbarium labels 
as a clump of tall, suckering stems growing among boulders in the streambed. 
The plant was evidently immature when Pollard first collected it, and he revis- 
ited it several times to get more mature material. The first three specimens (1955, 
1956, and 1958) are all sterile, and the 1956 label specifically describes the plants 
as “Adventive seedlings,” while the 1959 collection is the only fertile specimen 
in the series, and the label says: “Largest shoots, heretofore sterile, beginning to 
bear fruit.” The 1955 collection was annotated, apparently in Pollard’s hand- 
writing, as F. carica; the others were named “Ficus pseudo-carica Miq.?,” all three 
with the question mark. 

Morphological characters of these specimens are given in Table L. 


— 


RESULTS 


The Pollard collections from Cold Spring Canyon match F carica, not F palmata, 
in most of their characters (the thickness and dark coloration of the twigs, the 
large, deeply lobed leaves with obtuse or acute apices, strongly cordate bases, 
and shallowly crenate margins, and the turbinate fruit). Only the density of 
the indumentum on the twigs and petioles is in any way unusual for F carica, 
and this cl is rather variable in F carica. The indumentum is dense only 
in the earlier Cold Spring Canyon collections (1955 and 1956, made when the 
plants were quite immature); later collections, made as they approached and 
reached sexual maturity (1958 and 1959), show sparser indumentum, within 
the normal range of variation of F carica. The plant collected by Pollard and 
reported by several authors as F. pseudo-carica or F palmata is clearly F. carica; 
the unusual indumentum characters may be related to juvenility. Specimens of 
F. palmata have occasionally been grown horticulturally in the United States 
(WE Wight 4870, Franceschi nursery, Santa Barbara California, NA), but there 
is no evidence that the species has ever escaped from cultivation. It should be 
excluded from the adventive flora of North America. 


Morus nigra in the southwest 
Black mulberry (Morus nigra) is a small tree native to the mountains of south- 
west Asia (probably Iran and the Caucasus). It isa very old cultigen—mulberry 


Taste 1.Comparison of morphological characters of Asian Ficus palm 


reports of F palmata.The last column notes for each morphological character which species is the best match for the California specimen 


ata, cultivated F.carica,and California specimens forming the basis of the North American 


Ficus palmata 


Ficus carica 


Basis of CA F. palmata 
report 


Material coded 


Twig color 


diameter mm 
wig indumentum 
ee length mm 
Petiole indumentum 
Blade length mm 
Blade width mm 
Blade shape 
Blade lobing 
Blade/lobe margin 
Blade base 
Blade apex 


Fruit shape 


Fruit width mm 


31 sheets from India, Nepal, 
Pakistan, Saudi Arabia 
Yemen, Eritrea and Ethiopia 


tan or brown, sometimes, greenish, 


usually with a + ashy cast from the 
white indumentum 

1.5-4 

moderately to densely hirsute 
16-70 

sparsely to densely hirsute, 
seldom puberulent 

57-142 

50-140 

ovate to triangular 


usually unlobed, seldom 3(-5)- 
lobed to 0.4(-0.7) of length 
shallowly to strongly crenate- 
toothed or bluntly toothed 
broadly obtuse to truncate 

or shallowly cordate 
rounded, apiculate or 
acuminate 

globose, often from a linear 
stipe-like base 

9-15 mm 


ca. 70 sheets, cultivated 
worldwide 


brown, usually dark 


4-5 

glabrous or sparsely hirsute 
40-119 

glabrous or sparsely hirsute 


114-211 

129-210 

ovate to very broadly ovate 
in outline 

3-5-lobed 0.5-0.85 of length 
rarely shallowly lobed) 
crenate or irregularly wavy 


—a 


shallowly to deeply cordate, 
rarely broadly obtuse 

rounded or obtuse, ver 

rarely acute but never acuminate 
turbinate 


more than 15 mm 


4 Pollard collections, W 
fork Cold Spring a 
Santa Barbara Co. 
purplish brown 


sparsely to moderately hirsute 
110-125 
sparsely to moderately 


240-370 

ovate to very broadly 
ovate in outline 

5- se 0.75-0.8 of 


cordate 
obtuse or acute 
turbinate 


too immature to measure 


Best match for 
Pollard figs 


F.carica 


F.carica 
intermediate 
F.carica 

F. palmata 
F.carica 
F.carica 
either 
F.carica 
F.carica 
F.carica 


F.carica 


F.carica 


(L)@2 VaIs/9¥O'LIYS 


WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 773 


seeds are known from early archaeological sites in Mesopotamia and Egypt 
(Brothwell & Brothwell 1969)—and black mulberry is now widespread in cen- 
tral and western Asia and the Mediterranean basin. In the North American 
botanical literature, the name M. nigra has been confused since Small (1903, 
1933) misapplied it to dark-fruited forms of M. alba, an east Asian species that 
is widely naturalized in the United States (Wunderlin 1997). Wunderlin’s dis- 
cussion seems to suggest that the two taxa may be conspecific, but M. alba and 
M. nigra are actually quite distinct, differing (among other features) in their 
chromosome numbers, M. alba being diploid with 2n=28 whilea variety of poly- 
ploid or high aneuploid numbers, varying from 2n= 89-308, have been reported 
for M. nigra (Voltattorni 1947; Darlington & Wylie 1955; Hans 1972). 

Morus nigra was reported to escape occasionally in Beatty Townsite, Nye 
Co., Nevada, by Beatley (1976). This report has not been mentioned in subse- 
quent publications, including the Flora of North America Moraceae treatment 
(Wunderlin 1997). A check of Beatley’s hers showed that they are correctly 
determined and also revealed specimens of apparent waifs collected by Henry 
M. Pollard at a site in southern California (see below for specimen data). These 
collections indicate that black mulberry is able to reproduce as a waif at widely 
separated sites in the southwestern United States. It is only known asa waif in 
disturbed areas close to settlements, and there is no evidence that it can persist 
for long periods or disperse into intact natural vegetation. The species is not 
correctly described or keyed in any North American flora (except McMinn & 
Maino 1947, who included it as a cultivated species), soa brief description of M. 
nigra and an emended key for the region are provided. In order to enlarge the 
sample size and capture the normal range of variation in the Species the de 
scription below was supplemented with cultivated material from the south- 
western United States. Illustrations of M. nigra are provided by McMinn and 
Maino (1947, fig. 203) and Browicz (1982, fig. 1). 


KEY TO SPECIES OF MORUS IN THE WESTERN UNITED STATES 


1. Leaves reniform-triangular to almost circular, base deeply cordate. Styles 3-5 mm 
aieee densely hairy all over. eae purple-black or black M. nigra 
. Leaves ovate or triangular-ovate, base rounded, truncate, or shall rdate. Styles 
1-2 mm long, glabrous ioe with a few hairs). Syncarp red, adiele or white. 
2. Upper surface of leaf smooth, often very shiny, glabrous or with a few hairs on 
the main veins; underside glabrous except for scattered hairs on the major veins 
or in vein axils. Syncarp short-cylindrical to almost spherical, 0.6-1.8 X 0.5-0.7 


M. alba 


2. Upper surface of leaf dull, often scabrous, with hairs (very inconspicuous in M. 
rubra) evenly scattered over the blade; underside pubescent with hairs evenly 
scattered over the veins and blade. Syncarp various 
3. Trees to 20 m tall. Leaves ovate to subcordate or almost circular, 7-28 * 7-25 
cm, if lobed then lateral lobes abruptly acuminate. Syncarp short-cylindrical, 
1-1.9 X 0.6-0.8 cm M. rubra 


a 


774 BRIT.ORG/SIDA 22(1 


= 


3. Low shrubs to small trees 2-5 m tall. Leaves ovate or triangular-ovate, 2.8-8.3 
 1.6-5.3 cm, apex abruptly acuminate, if lobed then lateral lobes acute, ob- 
tuse, or occasionally rounded. Syncarp more or less spherical, 0.5-1.2 * 0.6- 
13cm M. microphylla 


Morus nigra L., Sp. Pl. 2:986. 1753. BLACK MULBERRY 


Small trees 3-5 m tall. Twigs brown, 2-4 mm thick, puberulent, pilose, or gla- 
brous; buds ovoid, 5-7 mim long. Petiole 1.2-3.0 cm long, pubescent with long 
soft hairs, at least ventrally. Blade reniform-triangular to almost circular, 
unlobed or rarely 3-lobed, 7-16 cm long and wide, (3-)5-veined from base; base 
deeply cordate, apex abruptly short-acuminate, margins dentate, teeth rounded- 
obtuse, 2-5 mm long; upper surface green, smooth or very weakly scabrous, gla- 
brous or with a few scattered hairs mostly confined to the major veins; under- 
side lighter green, soft-pubescent on veins and blade. Unisexual. Styles 3-5 mm 
long, densely hairy all over. Fruiting peduncle densely pubescent; syncarp short- 
cylindrical, 1.4-2.2 cm es 10-16 cm wide, purple-black or black. 


Specimens examined. U.S.A. NEV, e Co eee) escaped, in thicket with Screw-bean, 
Beatty townsite, eo ee USA. E.C’s Nevada Test Site, J. Beatle y 13438 (US). CALI- 
FORNIA, Ventura Co.: two small trees in low waste ground in dense brake of Rubus procerus, S of 
railroad and Fox St. drain, Ojai, origin unknown, location suggesting escape from cultivation, H.M. 
Pollard s.n., 21 Oct 1965 (NA); same locality, H.M. Pollard s.n., 20 Jun 1968 (NA). 


ACKNOWLEDGMENTS 
I would like to thank the curators at CAS, MO, UC and US for providing facili- 
ties for work at their herbaria, and Guy Nesom, Steve Boyd, Dieter Wilken, and 
Margriet Weatherwax for helpful comments. 


REFERENCES 


Beatey, J.C. 1976. Vascular plants of the Nevada Test Site and central-southern Nevada. 
Springfield, VA, National Technical Information Service, 1D-26881. 

BrotHWwett, D.and P. BrotHwelt. 1969. Food in antiquity. Ed. 2. Baltimore: Johns Hopkins Uni- 
versity Press. 

Browicz, K. 1982. Moraceae. Flora lranica fascicle 153. Graz: Akademische Druck- u. 
Verlagsanstalt. 

Conor, |.J. 1947. The fig. Waltham, Mass.: Chronica Botanica Co. 

DaruncTon, C.D, and A.P. Wyue.1955.Chromosome atlas of flowering plants. Ed. 2. London: 
Allen & Unwin Ltd. 

Friis, |. 1993. Moraceae. In: M. Thulin, ed. Flora of Somalia, vol. 2. London: Royal Botanic 
Gardens, Kew. Pp. 91-104. 

GuHaroor, A. 1985.Moraceae. Flora of Pakistan fascicle 171.Islamabad: Pakistan Agricultural 
Research Council; and Karachi: Dept. of Botany, University of Karachi. 

Hans, A.S. 1972. Cytomorphology of arborescent Moraceae. J. Arnold Arbor. 53:216-225. 

Hrusa, F., B. Ertrer, A. SAnbers, G. Lepric, and E. Dean. 2002. Catalogue of non-native vascular 


WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 775 


plants occurring spontaneously in California beyond those addressed in The Jepson 
manual—part |. Madrono 49:61-98. 

McCuntock, E. 1993. Moraceae. In: J.C. Hickman, ed. The Jepson manual: higher plants of 
California. Berkeley: University of California Press. Pp. 764-765. 

McMinn, H.E.and E. Maino. 1947. An illustrated manual of Pacific Coast trees. Ed. 2. Berkeley: 
University of California Press. 

Munz, PA. 1974.A flora of southern California. Berkeley: University of California Press. 

Smatt, JK. 1903. Flora of the southeastern United States. New York: Published by the 
author. 

Sma, J.K. 1933. Manual of the southeastern flora. New York: Published by the author. 

Smito, C.F. 1976. A flora of the Santa Barbara region, California. Santa Barbara: Santa 
Barbara Museum of Natural History. 

Smit, C.F. 1998. A flora of the Santa Barbara region, California. Ed. 2. Santa Barbara: Santa 
Barbara Botanic Garden. 

Voutattorni, S. 1947. Embriologia e cariologia di Morus nigra L. Annali Sperim. Agrar. N.S. 


Wunoertin, R.P. 1997. Moraceae. In: Flora of North America Editorial Committee, Flora of 
North America, volume 3. New York: Oxford University Press. Pp. 388-399. 


776 BRIT.ORG/SIDA 22(1) 


Book REVIEW 


CHET VAN Duzer. 2004. Floating Islands: A global bibliography with an Edition 
and Translation of G. C. Munz’s Exertacio academica de insulis natantibus 
(1711). USBN 0-9755424-0-0, hbk.). Cantor Press, Los Altos Hills, Califor- 
nia 94024, US.A. (Orders: www.cantorpress.com). $44.95, 400 pp., illus- 
trated, 6 1/8" x 9", 


To an average versed and educated reader the word floating island evokes association to Jules Verne 


and Jonathan Swift, to the description of explorations of swamps along the upper course of the Nile 

River and to the floating gardens of the Titicaca Lake. These do not seem entirely concrete objects 

anime in the aie life. This is we case in most countries without such islands. Floating Islands 
| review of 2500 years of publications on floating islands, cov- 


ering all plpeevile items from floating bogs to pumice islands and from bird nest rafts to floating 
air port 

o Duzer’s book consists of two main parts. The first (33 pp.) isa reprint of the full Latin text 
and English translation of Georgius Christophorus Munz’s rare 1711 Exercitatio acade mica de Insulis 
natantibus. Thus the basic work on the topic is easily accessible to the reader, both in its original 
form as well as its translation. Comprehensive and clear notes by Van Duzer follow (pp. 35-67) the 
Munz’s description. The second of the book is Global Bibliography on Floating Islands. On 280 
pages there are gathered more than 1800 references of books and articles in twenty languages treat- 
ing this subject. The entries are annotated and cross- ened and thet are e both h thematic and 


geographic indices. All aspects of floatin f floating 
islands, the causes of their buoyancy, ree role in the ecology of lakes aad wetlands, their flora and 
auna, their role in Bime eepeteal: st pate ana nee and methods for controlling and managing 
them. Worl ial {1 1 for agriculture, human habitation, wild- 
life habitat, and imp 
The book melndes the text al an sees aouapuuae Ww iui asiaied notes, of G.C. Munz’s rare 1711 
fl islands, E .as well as photographs of sev- 
eral floating aoa 

Van Duzer’s book is not only a complete (or one of the ae eiPiloarapiy on floating islands, 


f water ake: and floating ade in literature, myth, and legend. 


but also an indispensable source of literature for everyone { float- 
ing islands. Floating Islands succeeds in condensing a large body of historical ne and 
thought, in addition, it contains summaries of the recent research on floating wetlands. Essentially 


every feature of floating islands is s exarined—their formation, causes of their puCyaney, dominan 


1 } 
flora and fauna, and tt leir interactions with the wetland, lake, and river habitats with w hich they are 
1. Th ee ea 


associatec I d if ing islands, to their uses, including 


human habitation, and into classical inenanine and mythology. 

Van Duzer has not only provided a masterful treatment of the subject but has also provided an 
excellent sets fee ing ‘Islands should serve asa pasts for ne sc OlaeP for years tocome. 
Recommended fo ¢, botanical, herbarium Larger public librar- 


ies would be well cee to include it in their collection.—Gary Jennings, ee Research Insti- 
tute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


SIDA 22(1): 776. 2006 


EXOTIC PLANT INTRODUCTION IN KANSAS, 
TWO NEW SPECIES 


lralee Barnard 


962 300 Avenue 
Hope, Kansas 67451, U.S.A. 
findiralee@yahoo.com 


ABSTRACT 
Described in this paper are two new records for Kansas, Myagrum Bae coals Li <Buassleaceae) and 
eee thunbergii (DC.) Nakai (Fabaceae), with details about their lo and estab- 
lishmen 

RESUMEN 


Se aportan dos nuevas citas para cee ee de perfoliatum L. (Brassicaceae) y Lespedeza 
thunbergii (DC.) Nakai (Fabaceae), con de su introduccién local y establecimiento. 


Myagrum perfoliatum (Brassicaceae) was discovered in the ranch cattle pens 
south of the barn at Tallgrass Prairie National Preserve in Chase County, Kan- 
sas by Dillis Owen, National Park Service employee, in June of 2000. Plants were 
1 mtalland almost as wide. There were no leaves remaining on the plants when 
they were found. The distinctive fruits are club-shaped and 5-7 mm long. The 
primary associated species in the corrals at the time were Rumex, Amaranthus, 
and Conium maculatum. 

There are no other ds growing in Kansas that could be confused eas- 
ily with M. ee 


Voucher ited: KANSAS. Chase Co.: Tallgrass Prairie National Preserve, a few plants in the 
cattle pens near the barn, 10 Jun 2000, Dillis Owen s.n. (KSC 


Myagrum perfoliatum is native to Europe and western Asia. Exactly when it 
arrived in North America is uncertain, but it most likely came as a crop weed. 
This species is currently recorded from Texas, Oklahoma, and Ohio (USDA 
2005). In the Texas counties of Dallas, Delta, Denton, Frannin, Grayson, Johnson, 
and Rockwall it is described as abundant and spreading (Diggs et al. 1999). 

Cattle delivered to Tallgrass Prairie National Preserve are frequently 
shipped from Texas, so it is presumed that introduction of M. perfoliatum came 
from seed carried in the hooves, hair, or manure of livestock, or possibly on 
trucks when the animals were unloaded at the ranch corrals. 

When M. perfoliatum was first found at Tallgrass Prairie National Preserve, 
the plants were mature and had produced seeds. Subsequent search of the cor- 
rals revealed five locations, mostly along fences or along building perimeters, 
where M. perfoliatum was growing. The species persisted, and large, seed-pro- 


SIDA 22(1): 777 — 779, 2006 


778 BRIT.ORG/SIDA 22(1) 


ducing plants were collected again the following year in areas where they were 
missed by mowing. 

Intervening floristic surveys at Tallgrass Prairie National Preserve resulted 
in the discovery of no other populations on the ranch. Hand pulling, herbicide 
spraying, and mowing in the cattle pens have been effective in controlling this 
weedy annual. In 2004 and 2005 no M. perfoliatum was found in the corrals at 
Tallgrass Prairie National Preserve. Monitoring of the cattle pens for M. 
perfoliatum will continue. 

Lespedeza thunbergii (Fabaceae) is native to eastern Asia and has natural- 
ized in the eastern two-fifths of North America (Kartesz 1999, USDA 2005). In 
September 2005, L. thunbergii was found growing in sand prairie at Maxwell 
Wildlife Refuge in McPherson County, Kansas. The refuge is owned and oper- 
ated by Kansas Department of Wildlife and Parks. Manager, Cliff Peterson, re- 
calls that at one time a 3-row shelter belt ran east and west along the fence line, 
and L. thunbergii was planted in the north row of the shelter belt. The entire 
shelter belt and some of the fence has since been removed, but a population of 
L. thunbergii remains in ungrazed native prairie. 

During the same period that maintenance of the shelter belt was discon- 
tinued, refuge policy regarding prescribed burning changed. Beginning in 1978, 
fire was no longer totally suppressed as it had been for more than 30 years, and 
a third-year burning regime was initiated to reduce woody shrubs. This leads 
to interesting questions about the persistence of L. thunbergii at the refuge, since 
rather than inhibit shrub lespedezas, including L. thunbergii, fire favors them 
by encouraging seed germination (Maryland Cooperative Extension Service 
2005). 

Lespedeza thunbergii plants at the refuge are 1-18 m tall and grow in open, 
sandy soil among prairie grasses and forbs. They form a thicket approximately 
60 m? ina slight draw. Individuals are scattered a short distance from the main 
population. Multiple, semi-woody stems originate from the base of each plant. 
The rose-purple corolla is 13-15 mm long, leaflets are 2-3 times as long as wide 
and acute at the apex. The plants produced an abundance of seeds. 

The only other Lespedeza that might be mistaken for L. thunbergii is L. 
bicolor. Lespedeza bicolor has been found at only one or two locations in Kan- 
sas and differs by having a shorter corolla and rounder leaflets (Isely 1990). 


Voucher specimen cited: KANSAS. McPherson Co.:5 mi S, | mi W of Roxbury, SE of tower at Maxwell 
Wildlife Refuge in sandy prairie, 26 Sep 2005, Barnard 2086, C. Peterson (KSC). 

Lespedeza thunbergii is grown in the United States as an ornamental and is 
recommended for wildlife food and cover. The Maryland Cooperative Exten- 
sion Service has published the uses, establishment, and management of several 
shrub lespedeza species, including L. thunbergii (Maryland Cooperative Exten- 
sion Service 2005). The Southeast Exotic Pest Plant Council lists L. thunbergii 


BARNARD, FLORISTIC RECORDS FOR KANSAS 779 


as level three on the Kentucky invasive plant list. Level three is regarded as a 
“lesser threat” and described as a species that principally spreads and remains 
in disturbed corridors, not readily invading natural areas (SE-EPPC 2005). Con- 
cern about the invasive tendencies of related species, L. cuneata, in Kansas has 
persuaded refuge management in McPherson County to consider control or 
eradication of L. thunbergii. 


ACKNOWLEDGMENTS 


Specimens of both of these species are deposited at the Kansas State University 
Herbarium (KSC) in Manhattan, Kansas. I am grateful to Tallgrass Prairie Na- 
tional Preserve personnel for assistance, to Cliff Peterson for allowing permis- 
sion to collect at Maxwell Wildlife Refuge, and to Caleb Morse for input regard- 
ing KANU specimens. Thanks also to Craig Freeman and one anonymous 
reviewer for helpful suggestions. 


REFERENCES 


Dicas, G.M., B.L. Lipscome, and R.J. O’KeENNOoN. 1999. Shinners & Mahler’s illustrated flora of 
north central Texas. Biological Research Institute of Texas, Fort Worth. 

Isety, D. 1990. Vascular flora of the southeastern United States. Vol.3, Part 2 Leguminosae 
(Fabaceae). University of North Carolina Press, Chapel Hill. 

Kartesz, J.1.1999.A synonymized checklist and atlas with biological attributes for the vas- 
cular flora of the United States, Canada, and Greenland. First Edition.|In:Kartesz, J.T.,and 
C.A. Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina Bo- 
tanical Garden, Chapel Hill. 

MaryYLAND Cooperative Extension Service. 2005. FS-759-Wildlife plantings food and cover 
plantings shrub lespedezas. (http://www.agnr.umd.edu/MCE/index.cfm). University of 
Maryland, College of Agriculture and Natural Resources, College Park. 

SE-EPPC. 2005. Kentucky exotic plant list, version 3.0 (http://www.se-eppc.org). Bugwood 
Network The University of Georgia, Warnell School of Forest Resources and College of 
Agricultural and Environmental Sciences, Dept. of Entomology, Athens. 

USDA. NRCS. 2005.The PLANTS Database, version 3.5 (http://plants.usda.gov). Data com- 
piled from various sources by Mark Skinner. National Plant Data Center, Baton Rouge, 
LA 70874-4490 USA. 


780 BRIT.ORG/SIDA 22(1) 


Book NOTICE 


PAMELA A. Matson, ASHOK GADGIL, and DANIEL M.KAMMEN (eds). 2005. Annual Re- 
view of Environment and Resources: Volume 30, 2005. (ISBN 0-824 3-2330- 
O, hbk; ISSN 1543-5938). Annual Reviews Inc., 4139 El Camino Way, P.O. 
Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, 
800-523-8635, 650-493-4400, 650-424-0910 fax). $191.00, 648 pp., 6" x 9". 


The A IR FE. dR 


Th ol is an up-to-date review about changing 
environmental issues and the impact on the environment Om soc use Solas 
global resources.Contents of Volume 30 of Annual Review of E es 


I. Earth’s Life Support Systems 
Regional Atmospheric Pollution and Transboundary Air Quality Management 
Wetland Resources: Status, Trends, Ecosystem Services, and Restorability 
Feedback in the Plant-Soil System 

Il. Human Use of the Environment and Resources 


Productive Uses of Energy for Rural Development 


Private Sector Participation in the Water and Sanitation Sector 
Aquaculture and Ocean Resources: Raising Tigers of the Sea 
The Role of Protected Areas in Conserving Biodiversity and Sustaining Local Livelihoods 
Ill. Management and Human Dimensions 
Economics of Pollution Trading for SO, and NO, 
How Enyionmencal Health Risks Change with Development: The Epidemiologic and Environ- 
mental Transitions Revisite 
eel Value 
Righteous Oil? tone ene the Oil Complex, and Corporate Social Responsibility 
Archaeology and Global Change: The Holocene Record 


IV. Emexgmne Integrative Themes 


Adapt f Social-Ecological Systems 
Indexes 
Subject Index 
ulative Index of Contributing Authors, Volumes 21-30 
Cumulative Index of Chapter Titles, Volumes 21-30 
Errata 
An online log of corrections to Annual Review of Environment and Resources chapters may be 
found at http://environ.annualreviews.org. 


SIDA 22(1): 780. 2006 


NEW AND DISJUNCT RECORDS OF ELEOCHARIS LIESNERI 
(CYPERACEAE) FROM SOUTH AMERICA 


David J.Rosen 
S.M. macy mieraile 


{D 
Department GT ING! 1G 


as A&M ie iversity 
College Eo Texas 77843-2126, U.S.A. 


ABSTRACT 


Ext 


reported here for the first time from Brazil and Colombia. These newly discov- 
ered specimens represent the Le ene as pa the a) pe of the species, and suggest 
that this taxon might have a peri-A tion, key, and illustra- 
tion to aid in separation of E. liesneri from other South cee pede of Eleocharis subg. 


Limnochloa with which it has been confused are provided. 


RESUMEN 


Eleocharis liesneri se cita aqui por primera vez de Brasil y Colombia. Estos especimenes representan 


las unicas recolecciones conocidas aparte del tipo de la especie, y sugieren que este taxon pudiera 
tener una distribucion periamazonica. Se se una descripcion detallada, una clave, e ilustracion 
para ayudar en la separacion de - Henge de otras especies suramericanas de Eleocharis subg. 
Limnochloa con las que ha 


Eleocharis R. Br. is a cosmopolitan genus of about 200 species and 600 pub- 
lished names with a concentration of taxa in tropical America (Gonzalez- 
Elizondo & Tena-Flores 2000). Eleocharis subg. Limnochloa (P. Beauv. ex Lestib.) 
Torr. (= Eleocharis ser. Mutatae Syemeon)} is distinguished from other Eleocharis 
by acombination of usually cartil indurate, obscurely keeled, many- 
veined floral scales, coarse culms (often as thick as the spikelet) and biconvex 
(rarely trigonous) achenes usually conspicuously sculptured with polygonal 
cells (Gonzalez-Elizondo & Peterson 1997 

During examination of specimens for a systematic study of Eleocharis subg. 
Limnochloa, I discovered a collection from the State of Mato Grosso, Brazil, that 
was identified as EF. fistulosa (Poir.) Link, an illegitimate name for E. fistulosa 
(Poir.) Schult., which is currently treated asa synonym of E. acutangula (Roxb,) 
Schult. However, this specimen differs from E. acutangula in having terete rather 
than triquetrous culms, longer floral scales, larger achenes with more numer- 
ous longitudinal rows of epidermal cells, and belongs instead to Eleocharis 
liesneri S. Gonzalez & Reznicek. Additional collections of E. liesneri from near 
Sao Paulo, Brazil, and from Departamento de Antioquia, Colombia, were also 
discovered annotated as E. mutata (L.) Roem. & Schult and E. acutangula re- 
spectively. This species has not been recorded from Brazil or Colombia, being 


SIDA 22(1): 781 - 786. 2006 


782 BRIT.ORG/SIDA 22(1) 


previously known only from the type location in the highlands of SE Venezuela 
(Gonzalez-Elizondo & Reznicek 1996). The discovery of a specimen of E. liesneri 
from Sao Paulo, ca. 3500 km SE of the type location, suggests that this poorly 
known taxon might be found in suitable habitat throughout the Brazilian High- 
lands (Fig. l). The collection from Colombia suggests that E. liesneri has a peri- 
Amazonian distribution (personal communication, Socorro Gonzalez- 
Elizondo). However, additional collections and a better understanding of the 
natural history (particularly habitat requirements) of this wetland species are 
needed to confirm this distribution. The purpose of this paper is to report these 
disjunct collections, and provide a key and illustration that might assist others 
in discovering additional misidentified specimens of E. liesneri. 

Assuggested by Gonzalez-Elizondo and Reznicek (1996) Eleocharis liesneri 
is very near, and could be confused with E. obtusetrigona (Lindl. & Nees) Steud. 
(Fig. 2 d-f), an infrequently collected species belonging to subg. Limnochloa. 
Prior to the work of Gonzalez-Elizondo et al. (2002), E. obtusetrigona had been 
poorly defined, being reduced to synonymy under E. fistulosa by Svenson (1939); 
or treated as a variety of FE. mutata. (Clarke 1898) or E. fistulosa (Barros 1960). 
Both E. liesneriand E. obtusetrigona differ markedly from E. mutata in having 
usually terete rather than trigonous culms, larger achenes, a neck like constric- 
tion between the achene apex and tubercle, and larger floral scales. 

A key to separate Eleocharis liesneri from E. acutangula, E. mutata, and E. 
obtusetrigona, along with a description modified from Gonzalez-Elizondo and 
Reznicek (1996) and illustration are provided below. Achene lengths in the key 
include the tubercle. 


KEY TO AID IN IDENTIFICATION OF ELEOCHARIS LIESNERI 
IN SOUTH AMERICA 


1. Floral scales coarsely many veined (raised veins clearly discernable at 20); 
achene apex markedly constricted to a short neck, the neck sometimes obscured 
by the style base. 
2. Culms terete or obscurely 3-5-angled distally or rarely trigonous with the culm 
faces flat or slightly convex and the angles obtuse in some specimens of E. 
obtusetrigona 
3, Achenes 2.8-3.2 mm long, 1.3-1.9 mm wide with 23-26 longitudinal rows of 
transversely oblong cells; exposed portion of floral scales longer than wide 
E. liesneri (Fig. 2 a—c) 
3. Achenes (2.2)2.4—2.9(-3.1) mm long, 1.3-1.5(-1.6) mm wide with 11-18 lon- 


gitudinal rows of transversely reniform i posed portions of floral scales 
about as long as wide E. obtusetrigona (Fig. 2 d-f) 

2. Culms triquetrous distally, the culm f ale d the angles acute-winged 
E. acutangula 

|. Fl | fi many eined (raised veins not clea able at 20X);achene 


apex slightly constricted at the summit into a hard sae icici of the same 
texture and color as the achene E. mutata 


ROSEN, RECORDS OF ELEOCHARIS LIESNERI FROM SOUTH AMERICA 


% 


= Ci, Fi; 
~\ 


A 


a km 
0 250 500 1,000 


a + {| 0- 1000 
> ~ (> 1000 - 2500 


~ WM >2500 
ba Way =" - 


La | rd Pi pee ay Ret iri L COON Pp 
Fic. 1. Peri 


Eleocharis liesneri S. Gonzalez & Reznicek, Novon 6:361. 1996. (Fig. 2a—c). Tyre: 
NEZUELA. BOLIVAR: Gran Sabana, ca. 10 km SW of Karaurin Tepui at junction of Rio 
Karaurin and Rio Asadon (Rio Sanpa), gallery forest, 519°N, 61 03’W, 900-1000 m, riverbank, 

23 Apr 1988, R. Liesner 23698 (HOLOTYPE: MO} ISOTYPE: NY}). 


Perennial herb; roots coarse, fibrous, dark drab brown; rhizomes elongated, to 


784 BRIT.ORG/SIDA 22(1) 


ig & Nees) Steud. d—e. Detail of achene, perianth bristles, and 
it ! Inf 1 ~ a] t yt “39 7N0 MAN ie ie ae Cea ORME PRT 5 eee! | 


tubercle. f. Detail of spil 


7634 (C) by Neva Mikulicz. 


ROSEN, RECORDS OF ELEOCHARIS LIESNERI FROM SOUTH AMERICA 785 


ca. 1.8 mm thick, scales not seen, tubers absent; culms (when dry) terete or dis- 
tally obscurely 3-angled, 40-65 cm x 2-4.4 mm, rigid, internally spongy, with 
incomplete transverse septa, finely longitudinally striate, olive-brown; leaves 
reduced to sheaths, sheaths 2, apically notched, membranous, loose, friable, 
variable in color proximally from dark brown to maroon, distally cinnamon to 
rufous, apex acute; spikelet cylindric, narrowly-lanceolate, 1.6-2.4.cm X 3.9-4.5 
mm, acute, proximal scale empty, obtuse, amplexicaul, appearing asa continu- 
ation of the culm; floral scales appressed to weakly spreading, oblong, 5.5-6.2 
mm X 2.3-3.2 mm, distal and medial 0.2-0.5 mm hyaline-erose, cartilaginous, 
darkish centrally or subapical dark splotch, reddish-stramineous marginally, 
apex acute (the distally hyaline region obtuse-rounded if present), central area 
nearly flat, coarsely many veined, mid-vein not distinguishable abaxially, con- 
spicuous in adaxial view; perianth bristles 6-8, sub-equal, exceeding the tu- 
bercle, ascending, broad and strap-shaped proximally, coarsely retrorsely spinu- 
lose nearly to the base, the spinules coarse distally, becoming finer and more 
closely spaced proximally, stramineous; stamens 3; style trifid; achene bicon- 
vex, more or less obovoid to broadly obpyriform, 2.8-3.2 mm (including tubercle) 
x 13-19 mm, with 23-26 longitudinal rows of deeply concave transversely 
oblong polygonal cells visible through transparent periclinal layer on each 
achene face, dull, buff, maturing to shiny light brown, apex constricted to about 
0.4 times the width of the achene to a distinct neck (this sometimes obscured 
by the style base); tubercle dorsoventrally compressed, triangular, 0.5-1.2 mm 
x 0.7-0.9(-1) mm, dark brown. 
Specimen citations. BRAZIL. Mato Grosso do Sul: Central Brazilian Plateau, ca. 1-3 km W of Km 261 
Xavantina-Cachimbo road, in standing water in wet forest marsh, 12° 49'S, 51° 46’W, 19 Jan 1968, 
Philcox & Fereira 4124 (NY). Sau Paulo: Gesammelt in Sud-Brasilien um Sao Paulo, without date 
aaa eae? annotation a. C. B. Clarke dated 24 Apr 1906), Usteri s.n. (ZT). COLOMBIA. 
o. de Sonson, km 4 of road Sonson-La Union (52 km from La Union), 
dictated roadside 8 - 44'N, 75° 18 W, 2480 m, 04 Oct 1987, Zarucchi et al. 6240(MO). 


ACKNOWLEDGMENTS 


lam very grateful to the herbaria indicated for loan of specimens, to Socorro 
Gonzalez-Elizondo for advice, encouragement, and helpful comments, to Neva 
Mikulicz for preparing the excellent illustrations, and Tony Reznicek for re- 
viewing this manuscript. Thanks to Larry Brown (SBSC) for requesting loans 
of the type specimens. Financial support for research was provided by the Frank 
W. Gould Award for Graduate Student Research in Plant Systematics and the 
Dr. Harry Wayne Springfield Graduate Student Endowment. 


REFERENCES 


Barros, M. 1960.Las ciperaceas del estado de Santa Catalina. Sellowia 12:262. 
Ciarke, C.B. 1898. Cyperaceae. In: R. Chodat, Plantae Hasslerianae soit €numération des 


786 BRIT.ORG/SIDA 22(1) 


nlantes récoltées par le Dr.Emile Hassler, d’Aarau (Suisse) de 1885-1895 et déterminées 
par le Prof. Dr. R. Chodat avec l'aide de plusiers collaborateurs. Bull. Herb. Boissier 6, 
Appendix 1:19-22. 

GonzAez -Euizonpo, M.S.and A.A. Reznicek. 1996. New Eleocharis (Cyperaceae) from Venezu- 
ela. Novon 6:356-365. 

GONZALEZ-ELIZONDO, M.S.and P.M.Peterson.1997.A classification of and key to the supraspecific 
taxa in Eleocharis (Cyperaceae). Taxon 46:433-449. 

GonzAtez-Evizonpo, M.S.and J.A.Tena-Fiores. 2000. Eleocharis (Cyperaceae) in the New World. 
In: K.L.Wilson and D.A. Morrison, eds. Monocots: systematics and evolution.CSIRO Pub- 
lishing. Australia. Pp.637-643. 

GonzALez-EuIzoNnbo, M.S., M. GONZALEZ-EvIZONDO, and S.G. SmitH. 2002. Eleocharis obtusetrigona 
(Cyperaceae) new to North and Central America. Acta Botanica Mexicana 60:7-11. 
Svenson, H.K. 1939. Monographic studies in the genus Eleocharis V. Rhodora 41:13-19; 

95-104. 


NEW AND SIGNIFICANT REGORDS.OF 
VASCULAR PLANTS FOR FLORIDA AND FOR 
COLEIFE COUNTYAND LEE COUN FYE RORIDA 


George J.Wilder Martha R.McCombs 
Division of Ecological and Social Sciences 28260 Pine Haven Way, Unit 85 
Florida Gulf Coast University Bonita Springs, Florida 34135, U.S.A. 
10501 FGCU Boulevard South mecombsmarthar@aol.com 
Fort Myers, Florida 33965-6565, U.S.A. 
gwilder@fgcu.edu 
ABSTRACT 


Documented records are reported for 89 taxa (81 species, 7 varieties, and one hybrid) from Collier 
County (31 taxa) and from Lee County (71 taxa), Florida (102 records altogether). A newly discovered 
hybrid, Carex gigantea x Carex lupuliformis, is discussed. 


Key Worps: Florida, native species, alien species, new records, hybrid 


RESUMEN 


Se realizan las citas documentadas de 89 taxa (81 especies, 7 variedades, y un hibrido) de Collier 


Sony iO taxa) y de Lee County (71 taxa), Florida (102 citas en total). Se discute un hibrido 
, Carex gigantea X Carex lupuliformis. 


INTRODUCTION 


Herein we present new documented records ont ian of vascular plants from 
Collier County and/or Lee County, Florida, and wv discuss a Carex hybrid 
here reported for the first time in the literature. 

Collier County and Lee County occur within the southwestern portion of 
the Florida Peninsula and border one another from south to north, respectively. 
These counties encompass 2,026 and 804 square miles, respectively (Southwest 
Florida Regional Planning Council 2002). They merit especial attention for five 
reasons. 

(1) Gann et al. (2002), who grouped Collier and Lee counties among South 
Florida’s counties, characterized South Florida as “.. one of the most biologi- 
cally diverse regions in North America.” They reported over 2,200 species of 
native and naturalized plants from South Florida, including over 1,400 native 
species. By contrast, Wunderlin and Hansen (2003) recognized less than twice 
as many native and nonnative taxa (species, infraspecific taxa, and hybrids) 
for Florida’s 67 counties, overall (.e., 4,145 taxa). Wunderlin and Hansen (2004) 
listed the following numbers of taxa for the ten South Florida counties recog- 
nized by Gann et al. (2002): Broward Co., 1001; Charlotte Co., 709; Collier Co., 
1194; Glades Co., 485, Hendry Co., +93; Lee Co.,, 1172; Martin Co., 988; Miami- 


SIDA 22(1): 787 -— 799. 2006 


788 BRIT.ORG/SIDA 22(1) 


Dade Co., 1655; Monroe Co., 575, 776 (the mainland portion of Monroe Co. and 
the Florida Keys, respectively); and Palm Beach Co., 1036. 

(2) South Florida extends into the subtropics, thus exhibiting impressive 
overlap between typically temperate, and characteristically tropical vegetation. 
Coordinate with their subtropical milieu, at all times of year Collier and Lee 
counties exhibit particular species in flower. 

(3) Both counties manifest intense residential and commercial develop- 
ment. Within extensive areas their floras are being decimated, and increasing 
numbers of taxa face extirpation or rarity. The Southwest Florida Regional Plan- 
ning Council (2002) defined Southwest Florida as consisting of six counties, 
including Collier and Lee counties. From 1950 to 2000, Southwest Florida’s share 
of total U.S. population increased over nine-fold, from 0.047% to 0.429%. Simi- 
larly, from 1950 to 2000 Southwest Florida’s share of Florida’s population in- 
creased nearly three-fold, from 2.571% to 7.550%, the latter increase transpired 
despite the circumstance that Florida’s population growth exceeded that of the 
U.S., overall. 

(4) Together with Hawaii, Florida ranks among the two states “..with the 
most severe nonnative species problems...” (Simberloff 1997). One-third of the 
South Florida flora consists of escaped, nonnative plants (Gann et al. 2002). 
Simberloff (1997) stated that two primary factors predispose Florida, particu- 
larly southern Florida, to invasion by, and damage from nonnative species: (a) 
“destruction and disturbance of native habitats and their replacement by novel 
habitats...” and (pb) *..the geographic features of tropicality, insularity, and the 
great expanse of aquatic habitats.” Simberloff (1997) considered the problem of 
nonnatives to be aggravated by the considerable tourism and transportation 
into Florida. 

(5) No flora has been published for Collier County or Lee County. 


“ 


Climate 

Annually, Collier and Lee counties exhibit a warm rainy season and cooler dry 
season. During the dry season shallow, rain-fed pools dry up and the land ap- 
pears drier, overall. 

Liudahl et al. 1998) cited data gathered at Fort Myers (Lee Co.)—20 miles 
north of Collier County—to characterize the climate of Collier County. Thus, 
those data represent both counties. They indicated annual average tempera- 
ture of nearly 74° F and temperatures ranging from an average within the low 
sixties in January to the low eighties during summer. Average daily maximum 
temperatures vary from 74.7° F January) to 91.1° F (August); corresponding 
daily minimum temperatures for the two months are 53.6° F and 74.1° F 

Frost develops few times, yearly (Liudahl et al. 1998). Collier and Lee coun- 
ties each exhibit USDA plant hardiness zone nos. 10a and 10b, and Lee County 
also exhibits zone Ob ( th ESE ZOTIES represent average ann ual m inim umtem pera- 
tures of 30°-35" F 35°-40° F and 25°-30° F respectively; Anonymous 2005). 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 789 


Average annual rainfall exceeds 50 inches. Approximately, two-thirds of 
rain falls from June to September, when average rainfall is nearly eight inches 
per month. Monthly average rainfall is less than two inches from November 
through January and is a little over two inches from February through April 
(Liudahl et al. 1998). 


Geology and soils 

Abutting the Gulf of Mexico and belonging to the Atlantic Coastal Plain, both 
counties exhibit low, flat terrain. They are part of a larger, unnamed, primarily 
southern and peripheral region of Florida exhibiting maximum elevation of 
70 feet above sea level (Schmidt 1997). Physiographic regions prominent within 
Collier and Lee counties are the Big Cypress Spur, Caloosahatchee Valley, Gulf 
Barrier Chain and Gulf Coastal Lagoons, Immokalee Rise, Reticulate Coastal 
Swamps, and the Southwestern Slope (Liudahl et al. 1998; Schmidt 1997). Within 
each county, outcrops and shallow subcrop rocks are either Pliocene or Pleis- 
tocene-Holocene in age (Randazzo & Jones 1997). 

Henderson, (1984) and Liudahlet al. (1998) surveyed the soils of both coun- 
ties. For Collier Co., Liudahl et al. (1998) grouped general soil map units under 
either of three main headings: (1) urban land and soils in urban areas, (2) soils 
on the flatwoods and hammocks and in sloughs, and (3) soils on prairies and in 
swamps and freshwater marshes. For Lee Co., Henderson (1984) grouped major 
units similarly, but under four main headings: (1) soils of the manmade areas, 
(2) soils of the flatwoods and sloughs, (3) soils of the swamps and sloughs, and 
(4) soils of the tidal areas and barrier islands. 


Ecosystems and land-use history 

Despite their minimal ranges of elevations, both counties, collectively, exhibit 
diverse ecosystems, e.g., pine flatwoods, scrub, diverse kinds of hammocks, 
swamps, freshwater marshes, saltwater marshes, mangrove forests, and ruderal 
sites (Myers & Ewel 1990; Taylor 1998; G. Wilder, pers. obs.). 

In addition to development, land use in Collier and Lee counties has en- 
tailed these extensive human-induced modifications: construction of drainage 
canals and of ditches for mosquito control; elimination of, and various alter- 
ations of wetlands; cattle ranching; row-crop agriculture; and lumbering. Both 
counties also maintain major preserves or portions of preserves: Big Cypress 
National Preserve, Collier Seminole State Park, Corkscrew Regional Ecosystem 
Watershed, Corkscrew Swamp Sanctuary, Fakahatchee Strand State Preserve, 
and Everglades National Park. Furthermore, Everglades restoration, currently 
underway partly within Collier and Lee counties, ranks among the largest of 
efforts for ecosystem restoration, worldwide. 


MATERIALS AND METHODS 


We made most collections between and including 2002 and 2005, however, we 
collected Limnobium spongia in 1994 and Leptochloa nealleyi, Paspalum 


790 BRIT.ORG/SIDA 22(1) 


distichum, and Scirpus pungens in 1998; Appendix). All collections but one were 
prepared as dried herbarium specimens; Landoltia punctata was fixed in aque- 
ous formalin-propionic acid-ethanol solution and stored in aqueous glycerin- 
ethanol solution. Specimens cited are deposited in the Herbarium of Southwest- 
ern Florida (SWF), housed at Florida Gulf Coast University (Fort Myers, FL). 
We characterize taxa as native, alien, and endemic, according to Wunderlin and 
Hansen (2003, 2004). Nomenclature for species and infraspecific taxa follows 
Wunderlin and Hansen (2003), but for several taxa we also provide commonly 
used synonyms. Family circumscriptions and common names also follow 
Wunderlin and Hansen (2003), except for Lemnaceae, Nyssaceae, and Asclepiada- 
ceae, which we keep separate from Araceae, Cornaceae, and Apocynaceae, re- 


spectively. 

The Atlas of Florida Vascular Plants Wunderlin @ Hansen 2004) was our 
principal source for determining whether taxa represented new county or state 
records. That database does not reference voucher specimens, however, 
Wunderlin (2002) specified that “Each distribution record on the atlas website 
is documented by an herbarium specimen or a reliable published source, such 
asa monograph or revision.” We also recognized documented plant records from 
Austin et al. 1990) and Gann et al. (2002). 

There exists a plethora of additional reports and inventories pertaining to 
the floras of Collier County and Lee County, which do not reference voucher 
specimens, which are unpublished, and/or which have insufficient circulation. 
Because of these shortcomings, we do not consider records cited within them 
as adequately established. As Wunderlin (2002) stated in regard to voucher 
specimens, if a “..species is not documented..there is no record that the plant 
ever existed...” 


RESULTS AND DISCUSSION 
Reported herein are 89 taxa (81 species, 7 varieties, and one hybrid) that repre- 
sent 31 new documented records for Collier Co. and 71 for Lee Co., Florida (02 
records altogether). The hybrid, Carex gigantea < Carex lupuliformis, is newly 
discovered and thus a new Florida record. 

Asclepias feayi is included here, although it was documented previously. 
That species was originally collected near Immokalee (Collier Co.) in 1967 (Gann 
etal. 2002). It is cited here, because those workers designated A. feayi as histori- 
cal in South Florida Gincluding Collier Co.), a designation implying that the 
species might have been extirpated there. 

Muss et al. (2003) published a plant-species list for the Big Cypress Na- 
tional Preserve, which includes portions of Collier, Miami-Dade, and Monroe 
counties. They listed nine species cited in this article: Cyperus involucratus, 
Dalbergia sissoo, Eleocharis flavescens, Ixora coccinea, Limnophila sessiliflora, 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 791 


Melilotus albus, Paspalum distichum, Reimarochloa oligostachya, and Senna 
alata, but did not indicate which, if any, records represented Collier County. 

The Institute for Regional Conservation (IRC) cites plant records for South 
Florida, from many sources (Gann et al. 2005). These records as presented by 
Gann etal.(2005) are undocumented and, therefore, not accepted here. Gann et 
al. (2005) cited four taxa that we report for Collier Co. (Eleocharis flavescens, 
Limnophila sessiliflora, Melilotus albus, Scirpus californicus) and eight taxa 
presently listed for Lee Co. (Asclepias feayi, Cirsium nuttallii, Commelina gambiae, 
Palafoxia integrifolia, Paspalidium geminatum, Paspalum conjugatum, Phyllan- 
thus amarus, Quercus chapmanii). 


Taxonomic data 

The 89 taxa represent pteridophytes (1 taxon; 1.1 % of taxa), monocotyledons 
(44 taxa; 49.4%), and dicotyledons (44 taxa; 49.4%). Numbers of families in 
each group are: | (pteridophytes), 10 (monocotyledons), and 24 (dicotyledons; 
Appendix). 

Families of monocotyledons with the most taxa are Poaceae (21 taxa) and 
Cyperaceae (15 taxa). Remaining monocotyledonous families each include one 
taxon. The 36 taxa of Poaceae and Cyperaceae, collectively, constitute 40.4 % of 
presently reported taxa. This disproportionately high percentage suggests that 
previous collectors within Collier and Lee counties documented these families 
insufficiently. Families of dicotyledons with the most taxa are Fabaceae (8), 
Asteraceae (5), Veronicaceae (5), Euphorbiaceae (4), Brassicaceae (2), and 
Urticaceae (2). Remaining dicotyledonous families each include one taxon. 


Native and Endemic Taxa 

Wunderlin and Hansen (2003) classified as native to Florida 48 (53.9%) of the 89 
taxa reported here (Appendix). They listed two of our taxa as endemic to Florida 
(Asclepias feayi and Linum carteri; Appendix). We consider Carex gigantea x 
Carex lupuliformis endemic, as well. In the Poaceae and Cyperaceae, the families 
most prominently represented in this study, 12 and 10 taxa are native, respec- 
tively (ie. 57.1% and 66.7% of taxa of these families). The present report of a large 
proportion (ca. one-half) of nonnative taxa accords fairly well with the consider- 
able proportion (ca. one-third) of such taxa indicated previously for South Florida 
(Gann et al. 2002) and for Florida, overall (Wunderlin & Hansen 2003). 


Rare Florida Taxa 

The eighty-nine records include a considerable number of rare taxa. Foremost, 
is Carex gigantea X Carex lupuliformis, first collected by George Wilder on April 
22, 2003. We found that hybrid solely at one station, in Six Mile Cypress Slough 
Preserve (Lee County). Two years ago, Dr. Anton Reznicek (MICH) annotated 
three herbarium specimens of this hybrid as “a beautiful intermediate collec- 
tion” (Wilder & McCombs [W & M] 17258, W & M 17259, and W & M 17260). 


792 BRIT.ORG/SIDA 22(1) 


Preliminary study revealed three differences between both parental spe- 
cies and the hybrid. () Perigynia exhibit achenes (both parental species), but 
are sterile (the hybrid); indeed, sterility isa common feature of hybrids, overall. 
(2) Relative to spike axes, perigynia extend at, or approximately at right angles 
(C. gigantea), are ascending (C. lupuliformis), and have generally intermediate 
orientations (the hybrid). (3) Spike diameters (excluding perigynium beaks) 
are narrow (C. gigantea), broad (C. lupuliformis), and intermediate (the hybrid). 

Cayouette and Catling (1992), in their comprehensive review of Carex hy- 
brids, referred to “Carex gigantea Rudge x lupulina Willd. @ lupuliformis 
Sartwell). Possible hybrid origin according to Correll and Johnston (1970).” That 
language is interpretable in various ways—e.g., to mean the existence of C. 
gigantea X C. lupuliformis, however, the latter interpretation is incorrect. Correll 
and Johnston (1970) (who construed C. lupuliformisasa form of Carex lupulina 
Muhl., rather than as a discrete species) actually said of C. lupuliformis, that “ 
..these plants seem to show the introgressive influence of C. lupulina by C. 
gigantea.” Thus, Correll and Johnston (1970) suggested hybrid character of C. 
lupuliformis, overall, not a definite C. gigantea/C. lupuliformis hybrid. 

Carex gigantea X Carex lupuliformis was not reported or recorded in Flora 
of North America (Reznicek 2002), in section Lupulinae of the genus Carex, 
nor was it indicated in floristic manuals listing both parental species (Fernald 
1950; Gleason 1968: Gleason & Cronquist 1991; Godfrey & Wooten 1979; 
Mohlenbrock 1999; Small 1933; Wunderlin & Hansen 2003). Some of these work- 
ers submerged, or apparently submerged Carex lupuliformis into Carex 
lupulind Mull. 

Leptochloa nealleyi and Kyllinga hyalina are also unusual in Florida. George 
Wilder collected L. nealleyi on Sanibel Island (Lee Co.), on Aug, 27,1998 (W & M 
9774). We know of one other Florida collection of L. nealleyi, from Miami-Dade 
County (Keith A. Bradley 2583, Aug. 19, 2004). Leptochloa nealleyi was listed by 
Wunderlin and Hansen (2004), but not by Wunderlin and Hansen (2003). 

We collected Kyllinga hyalina in Collier County and Lee County, where it 
isa lawn weed and grows in disturbed areas. Two years ago, Dr. Anton Reznice 
(MICH) annotated six of our specimens as “Cyperus hyalinus Vahl 2nd US. col- 
lection” (W & M 17261 to W & M 17266, all of Sept. 8, 2002). (Wunderlin & 
Hansen [2002] listed Cyperus hyalinus Vahl and Kyllinga hyalina (Vahl) T. 
Koyama as synonyms). 

Wunderlin and Hansen (2003) ranked 17 presently reported taxa as rare in 
Florida. We increase that number by two, by including Carex giantea X Carex 
lupuliformisand Leptochloa nealleyi (taxa which those workers did not consider), 
Based on data from Wunderlin and Hansen (2003) and on our assessments of 
these two taxa, the 89 taxa considered here are ranked for Florida, as follows: rare, 
19 taxa (21.3% of all 89 taxa); occasional, 48 taxa (53.9%); common, 4 taxa (4.5%): 
and frequent, 19 taxa (21.3%). Rare and occasional taxa predominate among our 


—— 


N 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 793 


collections—composing, collectively, three-quarters of taxa presently listed. Com- 
mon and frequent taxa represent, collectively, ca. one quarter of taxa. 

Wunderlin and Hansen (2003) listed as rare in Florida the following of our 
taxa: Acalypha arvensis, Acalypha setosa, Achyranthes aspera var. pubescens, 
Alysicarpus vaginalis, Callitriche pedunculosa, Chamaesyce lasiocarpa, 
Commelina gambiae, Dalbergia sissoo, Heteropogon contortus, Ixora coccinea, 
Kyllinga hyalina, Leptochloa virgata, Linum carteri, Paspalum nicorae, Senna 
alata, Spigelia anthelmia, and Zoysia tenuifolia. Five of the nineteen taxa des- 
ignated as rare by Wunderlin and Hansen (2003) and by ourselves, collectively, 
are native to Florida (Appendix). 

For Florida, Coile and Garland (2003) classified Linum carteri as endan- 
gered and Tillandsia variabilis[Tillandsia valenzuelana A. Rich.Jas threatened. 

Gann et al. (2002) considered as rare in South Florida eight species pres- 
ently listed. They indicated two of these species as extirpated (Lipocarpha 
maculata [“collected once in 1965 near Immokalee”], Scirpus californicus [“last 
collected in 1965 near South Bay in Palm Beach County”), two species as his- 
torical (Asclepias feayi [“last collected in 1967 near Immokalee”|, Reimarochloa 
oligostachya [“last collected in 1977"]), and four species as critically imperiled 
(Leptochloa virgata, Nyssa sylvatica, Rhynchospora baldwinii, Rhynchospora 
wrightiana). 

We documented Gann et al’s (2002) eight taxa from various localities: 
Lipocarpha maculata (Lee Co,, 2 localities), Scirpus californicus (Collier Co., | 
locality), Asclepias feayi (Collier Co., 1 locality; Lee Co., 6 localities), 
Reimarochloa oligostachya (Collier Co., 1 locality), Leptochloa virgata (Lee Co., 
3 localities), Nyssa sylvatica (Lee Co., 1 locality), Rhynchospora baldwinii (Col- 
lier Co., 1 locality; Lee Co., 1 locality), and Rhynchospora wrightiana (Collier 
Co., 1 locality). 


~— 


— 


Directions and extents of Range Extensions 

Data from Wunderlin and Hansen (2004) indicate that 24 of our taxa repre- 
sent range extensions southward in Florida: Acalypha setosa, Anthaenantia 
villosa, Callitriche pedunculosa, Chasmanthium nitidum, Cyperus lanceolatus, 
Dalea carnea var. albida, Dichanthelium strigosum var. leucoblepharis, 
Hemarthria altissima, Iris virginica, Lindernia dubia var. dubia, Poa annua, 
Monotropa uniflora, Nyssa sylvatica, Portulaca amilis, Rhynchospora baldwinii, 
Rhyncho-spora fil vou id, Rite vere aaa Run: obovatus, Scirpus 


— 


californicus, Solida sis var. scabra, Sp} _Stylosanthes 
biflora, Urochloa texana, and Urtica namdedy oides. 
Fewer within Florida are northward (Leptochloa virgata), 


westward (Gonce coccinea, Spigelia anthelmia), both southward and eastward 
(Sida santaramensis), and both northward and westward (Kyllinga hyalina, 
Leptochloa nealleyi). 


794 BRIT.ORG/SIDA 22(1) 


For each of our taxa except the Carex hybrid, we gauged the relative dis- 
tance between the county (or two counties, collectively) newly represented here 
and the nearest county indicated by Wunderlin and Hansen (2004). For 60 taxa 
the newly reported county (or counties, collectively) bordered one or more of 
the indicated counties. For 2] taxa one or two counties intervened between the 
newly reported county (or counties, collectively) and the nearest county. For 
five taxa the number of intervening counties was 3 or 4 (Iris virginica, 
Callitriche pedunculosa, Dalea carnea var. albida, Stylosanthes biflora, Sida 
santaremensis). For three taxa between 6 and 8 counties intervened 
(Dichanthelium strigosum var. leucoblepharis, Solidago canadensis var. scabra, 
Lindernia dubia var. dubia). 

Prognosis for presently discussed taxa 

Development and habitat destruction are explosive within Collier County and 
Lee County, and we anticipate accelerated destruction of taxa and habitat in 
these counties. Certain localities represented herein have already been destroyed, 
and the following taxa are now extirpated from one or more localities repre- 
sented in this paper: Asparagus aethiopicus, Cirsium nuttallii, Cyperus 
sphacelatus, Melilotus albus, Melilotus indicus, Paspalum conjugatum, Paspalum 
notdatum var. notatum, Penstemon multiflorus, Polygala verticillata, and 
Urochloa texana. 


APPENDIX 
The following list of species, varieties, and a hybrid represent new documented 
records for Collier County and Lee County, Florida. 

Data are presented in the following order after the Latin name of a species, 
hybrid, or variety: relevant synonym, if any (between brackets); common name; 
the Wilder and McCombs collection number(s) of voucher specimen(s); 
habitat(s) where the species was collected; Wunderlin and Hansen’s (2003) as- 
sessment of frequency of the taxon in Florida; and county(ies) for which the 
taxon is presently documented. For individual taxa documented for both Col- 
lier Co.and Lee Co., collection numbers are indicated in the order listed of these 
counties. * = alien to Florida; © = endemic to Florida. 


PTERIDOPHYTE 
THELYPTERIDACEAE Mariana Maiden Fern—22,158;damp forest; 
“Macrothelypteris torresiana (Gaudich.) Ching, occasional; Lee 
MONOCOTYLEDONS 
ALISMATACEAE ASPARAGACEAE 
ss ittaria latifolia Willd, Broadleaf Arrowhead— — *Asparagus aethiopicus L. [Asparagus sprengeri 
22697; damp soil; occasional: Lee Regel], Sprenger’s Asparagus-Fern— 20731: 


scrub; occasional: Collier 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 


BROMELIACEAE 
Tillandsia HaMaoys is Schltdl., 
Airplant vamp; 
COMMELINACEAE 
*Commelina gambiae C.B. Clarke, Gambian Day- 
flower—22522; highly insolated, disturbed 
land; edge of field; rare; Lee 


CYPERACEAE 

xX Carex gigantea Rudge X Carex lupuliformis 
Sartwell ex Dewey— 19688; swamp; rare; Le 

*Cyperus involucratus Rottb. [Cyperus alternifolius 
L.J, Umbrella Plant—18666; insolated land 
beneath power lines; occasional; Collier 

oe lanceolatus Poir.,Epiphytic Flatsedge— 

etland within insolated, cleared 

ne iaeoleted land bordering trail; occa- 
sional; Lee 

*Cyperus sphacelatus Rottb., Roadside 
Flatsedge—19715, 18686; bulldozed, 
insolated land; insolated lawn; occasional; 
Collier, Lee 

Eleocharis flavescens (Poir.) Urb., Yellow 
ae 20749; along/on shaded trail 

ugh damp lowland; frequent; Collier 

ie inga aie (Vahl) T. Koyama [Cyperus 
hyalinus Vahl], Peduncled Spikesedge—dis- 
: Nise land; lawn; 18498, 17265; rare; Collier, 


Leatherleaf 


ccasional: Lee 


bie nga ane Michx.,L sedge—22033; 
disturbed land; oa Lee 
*Kyllinga squamulata Thonn. Ex Vahl, Asian 


ee pine flatwoods; occa- 
nal; Lee 
ree maculata (Michx.) Torr, American 
Halfchaff Sedge—18030; insolated wetland 
beneath power lines; disturbed land (for- 
merly either a flatwoods or scrub);on trail; 
occasional; L 
Rhynchospora i dwinii A. Gray, Baldwin's 
Beaksedge—19780; wet portion of pine 
flatwoods; occasional; Collier 
Rhynchospora filifolia A. Gray, Threadleaf 
eaksedge— 19084; periphery of pond; 
Fe eee llier 
Rhynchospora grayi Kunth, Gray’s Beaksedge— 
17911; pine flatwoods; occasional; Lee 
Rhynchospora wrightiana Boeck., Wright's 
Beaksedge—19812;along firebreak through 
pine flatwoods; occasional; Collier 


C irp r aif 


(C.A.Mey.) Steud., Giant Bul- 
tush—16350. edge of pond; occasional; 
Collier 

Scirpus pungens Vahl, Threesquare Bulrush— 
9790, 19209; insolated wetland; roadside; fre- 
quent; Collier, Lee 


HYDROCHARITACEAE 

Limnobium spongia (Bosc) Rich.ex Steud., Ameri- 
can Spongeplant—7392; occasional: in ca- 
nal; Lee 

HYPOXIDACEAE 

Hypoxis curtissii Rose, Common Yellow Star- 
grass—20142; swamp; common; Lee 

IRIDACEAE 

Iris virginica L., Virginia Iris—18207; insolated 
depression along road; occasional; Lee 

LEMNACEAE 

ti eee . oe Les & D. J. Craw- 

d Du 


| 1] 
JIIAHOVY weter; fre- 


ope ne 


POACEAE 
Anthaenantia villosa ane) P. Beauv., Green 
Cilkevweral Loew cei J ye ie 


JIIKYStdl I 


i.e) 2; pine fi ' | 


Lee 

*Bothriochloa ischaemum (L.) Keng, King Ranch 
Bluestem—19118; pasture; occasional: Lee 

Cenchrus gracillimus Nash, Slender Sandbur— 
19178, 21335; along dirt road through bull- 
dozed land; strand vegetation by Gulf of 
Mexico; frequent; Collier, Lee 

Chasmanthium nitidum (Baldwin) Yates, Shiny 
Woodoats—22291; Hammock; occasional; 


Lee 

Dichanthelium strigosum var. leucoblepharis 
(Trin.) Freckmann—19364; edge of trail 
through open woodland; occasional; Lee 

*Hemarthria altissima (Poir.) Stapf & C. E. Hubb, 
Limpograss—20728; edge of pond; occa- 
sional; Collier 

Heteropogon contortus (L.) P. Beauv. ex Roem. & 
Schult., Tanglehead— 19467; pine flatwoods; 
along railroad tracks; rare; Lee 

*Leptochloa  nealleyi Vasey, Nealley’s 
Sprangletop—97/74; disturbed land on 
Sanibel Island; rare; Lee 

Leptochloa virgata (L.) P. Beauv., Tropical 
Sprangletop—22364; damp depression 
along road; rare; Lee 


Panicum anceps Michx., Beaked Panicu 
26; wet prairie; pine flatwoods; on ai 
common; Lee 
doe idium - vatum (Forssk.) Stapf, Poyelal 
palidium—22686; insolated wetlan 


fell e 
Paspalum conjugatum PJ. Bergius, Sour 
spalum—21578;| k- recently bull- 
dozed, ae insolated land; along dirt road; 
occasional; L 


sc dsc mL, ANeghesS: 9787; dis- 
and; occasional; Collier 
enh nicorae Parodi, Brunswickgrass— 
21583:pine flatwoods; rare; Lee 
*Paspalum notatum Flugge var. notatum, 
Bahiagrass—213/70; ao highly 
insolated land; occasional: Le 


oO 
\O 
nn 
ea 
tS 


*Poa annua L.,Ann baile Olas “19550 
lawn:shaded,b 1erj 


frequent; Collier, Lee 

Reimarochloa oligostachya (Munro ex Benth.) 
Hitchc., Florida Reimargrass—21136; 
meadow; occasional; Collier 

*Secale cereale L., Cultivated Rye—22847; bull- 
dozed, highly insolated land; occasional; Lee 

eat olis obtusata (Michx.) Scribn., Prairie 

Wedgescale— 19594; field; frequent; Lee 

*Urochloa texana (Buckley) R. D. Webster, Texas 
Signalgrass - 19633; bulldozed, highly 
insolated sane a Lee 

— tenuifolia Willd. Thiele, Manila 

emplegrass— 19638, ok disturbed land; 

a Lee 


DICOTYLEDONS 


ACANTHACEAE 

*Thunbergia fragrans Roxb., Whitelady—22253; 
disturbed hammock, edge of field; occa- 
sional; Lee 


AIZOACEAE 

Sesuvium maritimum (Walter) Britton et al., Slen- 
der Seapurslane—21161;insolated trail; oc- 
casional; Coll. 


AMARANTHACEAE 
bcc aspera var. pubescens (Maq.) C.C. 
evil's Horsewhip—21102; oak- 
hee within Caloosahatchee 
River Regional Park; rare; Lee 


BRIT.ORG/SIDA 22(1) 


ASCLEPIADACEAE 

= Asclepias feayi Chapm.ex A.Gray , Florida Milk- 
weed—1/78/74; scrub, on/along trails 
through pine flatwoods, grassy portion of 
pineland remnant, disturbed land (formerly, 
apparently pine flatwoods); occasional; Lee 


ASTERACEAE 

Cirsium nuttallii DC., Nuttall’s Lae ae 
lawns (including tree lawn); frequen 

*Cyanthillium cinereum (L.) H. Rob. ae ci- 
nerea (L.) Less.]—18454; along/on dirt road; 

naliColller 

ae ae (Nutt.) Torr. & A. Gray, 
Coastalplain Palafox— 22101; field; pineland; 
frequent; Lee 

Solidago canadensis L.var.scabra (Mubhl.ex Willd.) 
orr. & A, Gray—18432, 17315; median strip 
of | 75; along road; wet von shallow wa- 


ter; frequent; Collier, Le 

*Taraxacum officinale ee ex F. H. Wigg. [in- 
cluding Taraxacum laevigatum (Willd.) 
DC]—19682, 18787 (red-fruited and brown- 


fruited formas, respectively); occasional; Lee 


BRASSICACEAE 


*Coronopus (L.) Sm., Lesser 
winecress 20397; highl val insolated 
dirt nian ae occasional; Collier, Lee 


Descurainia pinnata (Walter) Britton, Western 
Tansymustard—20401; tree lawns along 
main roads; common; Lee 


CALLITRICHACEAE 
*Callitriche pecunes losa Nutt., Ua 
Wraterctan rt phere) 


depression situated ee sen land be- 
neath power lines; rare; Lee 
ERICACEAE 
uniflora L., Indianpipe—16699; 
ha portion of scrub; occasional; Collier 


py ne REIACERE 
ilypha arvensis pp., Threeseed Mercury - 
17225; disturbed land by creek; rare; Lee 

*Acalypha setosa A. Rich., Cuban Copperleaf— 
18821;shrub bed within supermarket park- 


g lot; rare; Lee 

*Chamaesyce lasiocarpa (Klotzsch) Arthur, Road- 
side Sandmat—19225: field; tree lawn; 
insolated wasteland;crack in pavement; rare; 
Lee 


in 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 


ee amarus Schumach. & Thonn., Gale- 
— 21947; insolated, recently bull- 
a land of median of road; occasional; 


FABACEAE 
*“Alysicarpus vaginalis (L.) DC., White Money- 
wort 9; insolated, dry substrate of 
median of road;land by intersection; rare;Lee 
“Dalbergia sissoo Roxb. ex DC., Indian Rose- 
wood—21427; along railroad tracks; rare; 
Collier 
Dalea carnea var.albida (Tort.& A.Gray) Barneby, 
Whitetassels— 17934; pine flatwoods; occa- 
sional; Lee 
*Melilotus albus Medik., White Sweetclover— 
16471; field; frequent; Collier 
*Melilotus indicus (L.) All, Indian Sweetclover— 
905; bulldozed land; insolated dirt 
pile; occasional; Collier, Lee 
Mimosa quadrivalvis L., Sensitive Brier—17883; 
field; frequent; Lee 
*Senna alata (L.) roxb., Candlestick Plant— 19040, 
yee cea land; by lines; con- 
n site; rare; Collier, 
ene biflora (L.) se et on 22524; bor- 
dering, and extending into ditch; occasional; 


FAGACEAE 

Quercus chapmanii Sarg., Chapman's Oak.— 
17198; scrub; pineland; edge of disturbed 
forest; frequent; Lee 

ITEACEAE 

tea virginica L., Virginia Willo 20580; forest by 
creek; common; Lee 

LINACEAE 

2 Linum carteri Small, Carter's Flax—16958 field; 
rare; Lee 

MALVACEAE 

ae cise Monteiro, Moth Fanpetals— 

1; edge of field; occasional; Collier 

MORACEAE 

Morus rubra L.,Red Mulberry—19757; hammock, 
disturbed land; frequent; Lee 

NYSSACEAE 

Nyssa sylvatica Marshall, Tupelo—20834; shore 

f pond; frequent; Lee 


i 


797 


POLYGALACEAE 
Po oe a verticillata L., Whorled Milkwort— 
,edge of field; occasional; Lee 


POLYGONACEAE 

*Rumex obovatus Danser, Tropical Dock-—16909, 
insolated land by canal; dried-up ditch; 
weedy sod; occasional; Lee 


PORTULACACEAE 

“Portulaca amilis Speg., Paraguayan Purslane— 
20769, 21705; edge of insolated, bulldozed 
land; lawn weed; frequent; Collier, Lee 


RUBIACEAE 
“Ixora coccinea L., Scarlet Jungleflame—21085; 
natural land; rare; Collier, Lee 


STRYCHNACEAE 

Spigelia anthelmia L., West Indian Pinkroot— 
21232; insolated gravel and bulldozed land; 
tree lawn; meadow; rare; Collier; Lee 


URTICACEAE 
*Pouzolzia zeylanica (L.) Benn., Pouzolz’s Bush— 
oe land along road; insolated 
mp; occasional; Lee 
ae ica cramer Pursh, Heartleaf Nettle — 
3; lawn weed; occasional; Lee 


SS 
NOS 
~S 


VERBENACEAE 


*Verbena brasiliensis Vell., Brazilian Vervain— 
19764; depression by intersection; dried-up, 
barren portion of lawn; occasional; Lee 


VERONICACEAE 

*Limnophila — sessiliflora Blume, 
Marshweed—21181;damp substrate along 
ee 

Lindernia dubia var. anagallidea (Michx.) 
Cooperr., ee False Pimpernel.— 
21196;damp trench through insolated, bull- 
dozed land (growing together with the fol- 
lowing variety); occasional; Lee 

Lindernia dubia (L.) Pennell var. dubia— 21199; 
damp trench through insolated, bulldozed 
land; occasional; Lee 

Penstemon multiflorus (Benth.) Chapm.ex Small, 
Manyflower Beardtongue—22283; scrub; 
field; frequent; Lee 

*Russelia CACC OTS ae 2 nel, 


sional; Collier 


798 BRIT.ORG/SIDA 22(1) 


ACKNOWLEDGMENTS 


We thank Edwin Everham III, Aswani Volety, and Florida Gulf Coast Univer- 
sity for providing space, equipment, and herbarium supplies; the Coccoloba 
Chapter of the Florida Native Plant Society for financial support; Roger Clark, 
John Yarbrough, and the Lee County Dept. of Parks and Recreation for permis- 
sion to collect specimens in County parks and conservation lands; Bruce 
Hansen, John Kunzer, and Richard Wunderlin for identifying/confirming our 
identifications of virtually all taxa considered here; Anton Reznicek for con- 
firming our identification of Carex gigantea x Carex lupuliformis and for iden- 
tifying Kyllinga hyalina; Anik Smith, Barbara Conelly, and Richard Workman 
for directing us to single localities for Panicum anceps, Achyranthes aspera var. 
pubescens, and Sagittaria latifolia, respectively; Edwin Everham Ill, George 
Gann, Mark Garland, Bruce Hansen, Richard Wunderlin, and an anonymous 
reviewer for making suggestions about the manuscript; and Jordana Bitner, Brian 
Gilbert, Kathleen Gonzalez-Sanz, Tristan Gonzalez-Sanz, Debbie Preston, Jean 
Roche, Graham Walker, and Jennifer White for assistance in curating the Her- 
barium of Southwestern Florida. 


REFERENCES 


Anonymous, 2005. Florida Landscape. http://www.wgcu.org/fllandscape/zonemap.html. 
Florida Gulf Coast University, Fort Myers. 

Austin, D.F., J.L. Jones, and B.C. Benner. 1990. Vascular plants of Fakahatchee Strand State 
Preserve. Florida Sci. 53:89-122. 

Cayouerte, J.-and PM. Carina. 1992. Hybridization in the genus Carex with special reference 
to North America. Bot. Rev. 58:35 1-438. 

Cote, N.C.and M.A. Gartanp. 2003. Notes on Florida's endangered and threatened plants. 
Bot. Contr. No. 38, 4" ed. Florida Dept. Agric. & Cons. Serv., Div. Plant Industry, Gainesville. 

Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- 
search Foundation, Renner 

Fernato, M.L. 1950. Gray's manual of botany, 8" ed. D. Van Nostrand Co., New York. 

Gann, G.D., K.A. Braotey, and S.W. Woopmansee. 2002. Rare plants of South Florida: their his- 
tory, conservation, and restoration. The Institute for Regional Conservation, Miami, FL. 

Gann, G.D., K.A. Braotey, S.W.Woopomansee, and J.L. Sante. 2005. The floristic inventory of South 

Iconservation.org. The Institute for Regional Con- 


Florida database. http://ww 
servation, Miami, FL. 

GLEASON, H.A. 1968. The new Britton and Brown illustrated flora of the northeastern United 
States and adjacent Canada. Vol. 1. Hafner Publishing Co,, NY. 

Gteason, H.A. and A. Cronauist. 1991, Manual of vascular plants of northeastern United 
States and adjacent Canada, 2" ed. The New York Botanical Garden, Bronx. 

Goorrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United 


a7 


WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 799 


States. Vol. 1. Univ. of Georgia Press, Athens. 

HENDERSON, W.G. Jr. 1984. Soil survey of Lee County, Florida. U.S. Dept. Agriculture Soil Con- 
servation Service. 

LIUDAHL, K.eT AL. 1998. Soil survey of Collier County area, Florida. U.S. Dept. Agriculture Natu- 
ral Resources Conservation Service. 

Mon ensrock, R.H. 1999. The illustrated flora of Illinois. Sedges: Carex. Southern Illinois Uni- 
versity Press, Carbondale. 

Muss, J.D., D.F. Austin, and J.R. Snyper. 2003. Plants of the Big Cypress National Preserve, 
Florida. J. Torrey Bot. Soc. 130:119-142. 

Myers, R.L. and J.J. Ewer (eds.). 1990. Ecosystems of Florida. University of Central Florida 
Press, Orlando. 

Ranpazzo, A.F.and D.S. Jones (eds.). 1997. The geology of Florida. University Press of Florida, 
Gainesville. 

Reznicek, A.A. 2002. Carex Linnaeus sect. Lupulinge Tuckerman ex J.Carey.In: Flora of North 
America Editorial Committee, ed., Flora of North America north of Mexico. Vol. 23. Ox- 
ford University Press, NY. 

Scumiot, W. 1997.Geomorphology and physiography of Florida. |In:A.F. Randazzo and DS. 
Jones, eds. The geology of Florida. University Press of Florida, Gainesville. 

Simpertorr, D. 1997. The biology of invasions. In: D. Simberloff, D.C. Schmitz, and T.C. Brown, 
eds., Strangers in paradise. Island Press, Washington, DC. 

Smatt, J.K. 1933.Manual of the southeastern flora. Published by the author, New York. 

SOUTHWEST FLORIDA REGIONAL PLANNING COUNCIL. 2002. Strategic Regional Policy Plan.Vol.1.South- 


west Florida Regional Planning Council, Fort Myers. 

Taytor, W.K. 1998. Florida wildflowers in their natural communities. University Press of 
Florida, Gainesville. 

WunNDERLIN, R.P.2002.Where's the proof? The importance of herbarium specimens. Palmetto. 
21:6, 8. 

WUNDERLIN, R.P. and B.F. Hansen. 2003. Guide to the vascular plants of Florida, 2"? ed. Univer- 
sity Press of Florida, Gainesville. 

WUNDERLIN, R.P.and B.F. Hansen. 2004. Atlas of Florida vascular plants. http://www.plantatlas. 
usf.edu/isb/. Institute for Systematic Botany, University of South Florida, Tampa 


800 BRIT.ORG/SIDA 22(1) 


Books RECEIVED 


Martua J. GRoom, Gary K. Merve, and C. RONALD CARROLL. 2005. Principles of Con- 
servation Biology, Third Edition. (SBN 0-87893-518-5, hbk.). Sinauer As- 
sociates, Inc., 23 Plumtree Road, Sunderland, MA 01375-0407; U.S.A. (Or- 
ders: orders@sinauer.com, wwwsinauer.com, 413-549-4300, 413-549-1118 
fax). $92.95, 699 pp., 369 illustrations, 8 1/2" x 11" 


From the publisher—*Principles of Conservation Biology, Third Edition, features a wholly revised or- 
aie oe eae ing analyses of different categories of threat and approaches to conservation. 
Coverage has been expanded to incorporate both terrestrial and marine conservation issues, and 


ee the US. and across the globe. 


INDERJIT CEd.). 2005. Invasive Plants: Ecological and Agricultural Aspects. (ISBN 
3-7643-7137-4; 978-3-7643-7137-1, hbk.). Birkhauser Verlag, PO. Box 133, 
CH-4010 Basel, SWITZERLAND. (Orders: wwwspringer.com). $159.00, 283 
pp., illustrations, 6 1/2" x 91/4", 


From the publisher.—The aim ol this book is to discuss fundamental questions of invasion ecology, 
such as why particular communities become more invasible than others, what the mechanisms of 
exclusion of native species by invaders are, and whether invasion can be predicted. 

In addition, agricultural practices influencing invasion, the environmental and economic costs 


of invasion as w el as ao ee management strategies are discussed. Readers will get a unique per 


spective on inv gy through employing general principles of ecology to plant invasions. 


SIDA 22(1): 800, 2006 


CAREX BICKNELLII (CYPERACEAE) NEW TO ARKANSAS 


Paul M.McKenzie C. Theo Witsell 
U.S. Fish and Wildlife Service Arkansas Natural Heritage Commission 
101 Park DeVille Dr, Suite A 1500 Tower Building 
Columbia, Missouri 65203-0057, U.S.A. 323 Center St. 
paul_mckenzie@fws.gov Little Rock, Arkansas 72201, U.S.A. 
theo@arkansasheritage.org 
Joe Woolbright 


Ozark Ecological Restorations Inc. 
04 Skilern 
Siloam Springs, Arkansas 72761, U.S.A. 
joewoolbright@cox-internet.com 


ABSTRACT 


We report the first record of Carex bicknellii Britt. tor Arkansas. The species was collected by the 
authors in 2003 and 2005 from Baker Prairie Natural Area, an unplowed tallgrass prairie remnant in 


Boone County, Arkansas. 


RESUMEN 
Se realiza | le Carex bicknellii Britt para Arkansas. | | lecto por los autores 
en 2003 y 2005 en Baker Prairie Natural Area | | Boone County, 
Arkansas. 


The range of Carex bicknellii Britt. extends from Maine and Manitoba south to 
Kansas and South Carolina, with the bulk of the distribution extending from 
Missouri and southeastern Kansas northward to southeastern Manitoba and 
the western end of Lake Erie (Rothrock & Reznicek 2001; Mastrogiuseppe et al. 
2002). Disjunct populations occur in Maine, western Kansas, South Carolina, 
and the eastern Lake Ontario Region (Rothrock & Reznicek 2001). Habitat for 
the species includes dry to mesic prairies, rock, sand or serpentine barrens, open 
woodlands, openings in mesic to dry upland forests and bluff ledges, dry sandy 
or rocky fields, thickets, and less commonly along margins of ponds and fens 
(Yatskievych 1999; Rothrock & Reznicek 2001; Mastrogiuseppe et al. 2002). 
Orzell and Bridges (1987) and Smith (1988) reported C. bicknellii (as C. bicknellii 
var. bicknellii) for Franklin County, Arkansas but the specimen (Barber 861, 
UARK), which was the basis for the reports, was subsequently annotated by 
A.A. Reznicek as Carex bicknellii Britt. var. opaca FJ. Herm. J. Gentry, UARK, in 
litt. 21 October 2005). Hyatt (1988), Rothrock and Reznicek (2001), and 
Mastrogi et al. (2002) did not include Arkansas within the range of Carex 
bicknel Hii (either as C. bicknellii or as C. bicknellii var. bicknellii). Rothrock and 


SIDA 22(1): 801 — 804. 2006 


802 BRIT.ORG/SIDA 22(1) 


Reznicek (2001) referenced all specimens within the C. bicknellii group col- 
lected in Arkansas through 2000 and determined that all prior reports of C. 
bicknellii (as C. bicknellii var. bicknellii) in Arkansas were based on other species. 

On 9 May 2005, the senior author discovered this species scattered in ir- 
regular colonies on dry mounds of Baker Prairie Natural Area,a 28 ha remnant 
of native tallgrass prairie in Boone County in north central Arkansas located 
near the western edge of the town of Harrison. On 10 May 2005, McKenzie and 
Witsell returned to the site to assess the overall distribution and abundance of 
the species on the area and to record plant associates. Carex bicknellii was asso- 
ciated with Baptisia bracteata Muhl. ex Elliott, Berlandiera texana DC, Carex 
bushii Mack., Ceanothus americanus L., Commandra umbellata (L.) Nutt., 
Dodecatheon meadii L., Euphorbia corollata L., Hypoxis hirsuta (L.) Cov, Luzula 
bulbosa (Alph. Wood) Rydb., Physalis sp., Schizachyrium scoparium (Michx.) 
Nash, Silphium integrifolium Michaux, S. laciniatum L., and Tradescantia 
ohiensis Raf. Plants were irregularly scattered over much of the drier portions 
of the prairie and extended to a few of the swales that drain the site. Carex 
bicknellii was initially collected on Baker Prairie in a wet swale by Witsell on ] 
June 2003 but the specimen was incorrectly identified as the closely related C. 
opaca (FJ. Herm.) PE. Rothrock & Reznicek. 

With the exception of a small (approximately + ha) former pasture along 
its eastern edge, Baker Prairie Natural Area is a high quality unplowed prairie 
remnant, and is jointly owned by the Arkansas Natural Heritage Commission 
and the Arkansas Field Office of The Nature Conservancy. It is the only rem- 
nant of what was once an approximately 2000 ha tallgrass prairie located west 
and south of present day downtown Harrison and is the largest known rem- 
nant of Ozark prairie that occurs on a chert substrate in Arkansas (Arkansas 
Natural Heritage Commission 2004). Baker Prairie Natural Area occurs on the 
Boone Formation of the Springfield Plateau which is characterized by lime- 
stone with embedded chert (Harper et al. 1981, Arkansas Natural Heritage Com- 
mission 2004). The Natural Area is underlain by Nixa and Noark very cherty 
silt loams which are deep, well-drained, moderately permeable soils that formed 
in residuum from cherty limestone (Harper et al. 1981; Arkansas Natural Heri- 
tage Commission 2004). Elevation on the natural area ranges from 354 to 378 
m above mean sea level on slopes of 3 to 20 % (Arkansas Natural Heritage Com- 
mission 2004). Much of the natural area is gently rolling with scattered “pimple” 
mounds (Arkansas Natural Heritage Commission 2004). Baker Prairie includes 
dry-mesic chert prairie, mesic chert prairie (restricted to swales that bisect the 
area), and non-native grassland (Arkansas Natural Heritage Commission 2004). 
Major threats to the prairie prior to state ownership and restoration included 
lack of prescribed fire, excessive haying, the introduction of tall fescue (Festuca 
arundinacea Schreber) for grazing livestock, development pressure from adja- 


and 


— 


MCKENZIE ET AL., CAREX BICKNELLII NEW TO ARKANSAS 803 


cent communities, and the encroachment of woody vegetation (Arkansas Natu- 
ral Heritage Commission 2004). Due to the prairie’ ownership and such 
current management practices as regular use of prescribed fire and the control 
of exotics, most historical threats have been eliminated. One possible threat to 
plant species inhabiting Baker Prairie, however, is the potential for reduced ge- 
netic fitness due to the geographic isolation of this prairie from other high qual- 
ity prairies in Arkansas and adjacent states. 

Following European settlement and prior to state ownership, haying and 
grazing were apparently the main management tools used to maintain the prai- 
rie (Arkansas Natural Heritage Commission 2004). Beginning in 1992, Baker 
Prairie Natural Area has been the site of aggressive management efforts that 
include the removal of encroaching woody vegetation and exotic species; seed 
collection and planting; brush hogging; herbicide treatment of Festuca 
arundinacea, Lespedeza cuneata (Dum.-Cours.) Don, Lonicera japonica Thunb,, 
Carduus nutans L., Albizia julibrissin Durazz, Daucus carota L., woody sprouts 
and stumps; and the use of prescribed fire (Arkansas Natural Heritage Com- 
mission 2004). These intensive management efforts have been successful in 
maintaining the rich botanical diversity of the area and restoring the ecologi- 
cal integrity of the prairie. In addition to Carex bicknellii, Baker Prairie pro- 
vides habitat for other rare plant species tracked as elements of conservation 
concern by the Arkansas Natural Heritage Commission. These include 
Antennaria neglecta E. Greene, Aster sericeus Vent., Calopogon oklahomensis 
D.H. Goldman, Carex gravida L. Bailey, Gentiana puberulenta J. Pringle, 
Nemastylis nuttallii Picker ex R. Foster, Silene regia Sims, Trillium pusillum 
Michaux var. ozarkanum (EJ. Palmer & Steyerm.) Steyerm., Viola pedatifida G. 
Don, and Zizia aptera (A. Gray) Fern.. Rare animals of conservation concern at 
the site include ornate box turtle [Terrapene ornata ornata (Agassiz)], grass- 
hopper sparrow [Ammodramus savannarum (Gmelin)], willow flycatcher 
[Empidonax traillii (Audubon)], and prairie mole cricket (Gryllotapa major 
Saussure). 


Voucher Spec imens. U.S.A. ARKANSAS. Boone Co.: Ar} N l Heritage Commission and TNC’s 
Baker Prairie Natural Area in western Harrison, ca. 0.4 km $ of the inter. of Industrial Dr. and Goblin 
Dr; TI8N,R20WS5 SW1/4NE1/4N W1/4; Gaither 7.5' Quad.; N 360° 14'40.2", W 93° 08'01.6" (Datum = 
NAD83 Zone 15); infrequent in scattered colonies on dry mounds of native prairie; in association 
with Baptisia bracteata, Berlandiera texana, Carex bushii, Ceanothus americanus, Commandra 
umbellata, Euphorbia corollata, Hypoxis hirsuta, Luzula bulbosa, Physalis sp., Schizachyrium 
scoparium, Silphium integrifolium, S. laciniatum, and Tradescantia ohiensis,; 9 May 2005 Paul M. 
McKenzie 2187 (MO, MICH, UMO); 10 May 2005 Theo Witsell 05-0330 (anhe-Arkansas Natural Heri- 
tage Commission, UARK, peh-pers. herb. of Philip E. Hyatt); 1 Jun 2003 Theo Witsell 03-0437 (anhc). 


Botanical nomenclature listed herein follows Yatskievych and Turner (1990) 
except for Calypogon oklahomensis that follows Yatskievych (1999), and Carex 
bicknellii and C. opaca that follow Rothrock and Reznicek (2001). 


BRIT.ORG/SIDA 22(1) 


ACKNOWLEDGMENTS 
We are grateful to Anton Reznicek of the ey of Michigan Herbarium, 
Ann Arbor, Michigan, andanan ny for their assistance with this 


report. 


REFERENCES 


ARKANSAS NaturAL Heritace Commission. 2004. Baker Prairie Natural Area management plan. 
Arkansas Natural Heritage Commission. Little Rock. 

Harper, M.D., D.H. Fowckes, and D.A. Howard. 1981. Soil survey of Boone County, Arkansas. 
U.S. Dept. of Agric. Soil Conservation Service, in cooperation with the Arkansas Agri- 
cultural Experiment Station. 

Hyart, PE. 1998. Arkansas Carex (Cyperaceae): a briefly annotated list. Sida 18:535-554. 

Mastroaiusepre, J., PE. RoTHROCK, A.C. Digste, and A.A. Reznicek. 2002. Carex Linneaus sect. Ovales 
Kunth, Enum. PI. 2:394. 1837.|n: PW. Ball, K. Gandhi, R.W. Kiger, D. Murray, J.L.Zarucchi, A.A. 
Reznicek, J.L.Strother, eds.Flora of North America North of Mexico, vol.23 Magnoliophyta: 
Commelinidae: Cyperaceae. Oxford Univ. Press, NY. Pp. 332-378. 

Orzett, S.L.and E.L.Brioces. 1987.Further additions and noteworthy collections in the flora 
of Arkansas, with historical, ecological, and phyteographical notes. Phytologia 64: 
81-144 

RorHrock, PE.and A.A.Reznicek. 2001.The taxonomy of the Carex bicknellii group (Cyperaceae) 
and new species for Central North America. Novon 11:205-228. 

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Second Ed. 
Published by the author. Fayetteville, AR. 

YATSKIEVYCH, G. 1999. Steyermark’s flora of Missouri- Vol. 1- Revised Ed. Missouri Depart- 
ment of Conservation, Jefferson City, MO. 

YatskievycH, G. and J. Turner. 1990. Catalogue of the flora of Missouri. Missouri Bot. Gard. 
Monogr. Syst. Bot. No. 37. Braun-Brumfield, Inc, Ann Arbor, MI. 


GREEN FRINGED ORCHID (PLA TANTHERA LACERA, 
ORCHIDACEAE) IN SOUTHERN LOUISIANA 


Charles Allen, Sara Thames, 
ward Anderson lll, 


Bill Newton Ill,and Rhonda Hampton George Fisher 
Colorado State Univ., Fort Polk 509 Bon Ami 
1645 23° Street DeRidder, Louisiana 70634, U.S.A. 


Fort Polk, Louisiana 71459, U.S.A. 


ABSTRACT 


A new population of Platanthera lacera is reported for Vernon Parish, southern Louisiana. 


RESUMEN 


Se cita una nueva poblacion de Platanthera lacera de Vernon Parish, Sur de Louisiana. 


Green fringed orchid (Platanthera lacera (Michx.) G. Don) was reported for Loui- 
siana as early as 1852 by Riddell (1852). A specimen was collected in Shreveport 
in Caddo Parish by MacRoberts in 1976 (MacRoberts 1977). Apparently, this 
specimen was misidentified and not included in the orchid flora of Louisiana 
by Prigeon and Urbatsch (1977) and excluded by Thomas and Allen (1993). This 
species was reinstated to the Louisiana flora by MacRoberts and MacRoberts 
(1998): the authors also stated that the site for the collection in Caddo Parish 
was now a housing development. Green fringed orchid is reported from all of 
the eastern United States except for Florida (USDA, NRCS 2005). In Texas, this 
orchid is reported only from Bowie County (Correll 1947; Liggio & Liggio 1999). 

The habitats reported for this orchid include open sedge swamps and 
marshes, bogs, meadows, and glades of open woods, swampy woods and wet or 
occasionally dry open fields and prairies, and in thickets (Correll 1978; Liggio 
& Liggio 1999; Radlord et al. 1968). In Flora of North America Editorial Com- 
mittee (2002), this sp is reported from sphagnum bogs, alluvial and swamp 
forests, stream banks, riparian meadows, sand flats, moist and seeping slopes, 
prairies, roadside banks, ditches, old fields, and borrow pits. According to Liggio 
and Liggio (1999), this orchid has a preference for acidic soils but is indifferent 
to varying conditions of moisture, sunlight, and shade. 

On May 6, 2005, a new population of green fringed orchid was discovered 
in Vernon Parish, Louisiana which is ca 150 miles south of the Caddo Parish 
location and apparently the southernmost location in the United States. The 
collection data are: 


SIDA 22(1): 805 — 809. 2006 


806 BRIT.ORG/SIDA 22(1 


= 


Voucher specimen: LOUISIANA. Vernon Parish: along edge of mowed hunting trail, edge of pasture 
and beech/white oak forest in Section 14 T2S R7W, GPS N30.942 W93.053, ca. 1/4 mi SW of Talbert- 
Pierson Cemetery (Pine Grove Church) and ca. 5 mi NW of Sugartown in drainage of unnamed creek 
that drains into Whiskey Chitto Creek, 24 May 2005, Allen et al 19415, (BRIT, Fort Polk Herb., ULM). 


The habitat varied from an open mowed trail to the dense edge of the beech/ 
white oak forest of the creek edge and from moderately well drained to moist 
soil. The soil type is Gore which is a very fine sandy loam with 5 to 12 percent 
slopes (Soil Survey Staff 2003). The elevation in the area ranges from 150 to 160 
feet. The plants had just begun to flower on May 6, seemed to peak around May 
20, and just a very few flowers were noted on June 11, 2005. 

During May 2005, the site was revisited several times and 35 clumps of the 
orchid were noted; most clumps had a single flowering stem but a few had two 
flowering stems. The area surrounding each of the 35 clumps was examined 
and the five nearest individuals were identified including herbaceous, woody 
vine, shrub (woody non-vines shorter than six feet), shrubs/saplings (woody 
non-vines taller than six feet and five inch or less dbh), and trees (woody non- 
vines taller than six feet and larger than 5 inch dbh) species (Table 1). The per- 
cent of each species out of the 175 total individuals (five times 35) for each cat- 
egory is listed in Table |; for the herbaceous and woody vines only one percent 
is reported for each species and for the woody non-vine species, the first per- 
cent listed is the shrub, the second is the shrubs/saplings, and third is the trees. 
The surrounding area was examined for a distance of twenty feet and all asso- 
ciated species were identified and listed in Table 1. A total of 124 species from 
90 genera and 56 families were identified as a nearest individual or associated 
species. Densiometer readings were taken at each clump and the average cover 
percent was 61.04% and ranged from 43.03% to 75.00%. 

The most common occurring nearest herbaceous species was 
Chasmanthium sessiliflorum (Poir.) Yates (17.71%) and was followed b 
Dichanthelium dichotomum (L.) Gould (12.57%), Sanicula canadensis L. (8.57%), 
Allium canaddense L. (8.00%), and Dichanthelium laxiflorum (Lam.) Gould 
(5.71%). The woody vine species that was most commonly nearest the orchid 
was loxicodendron radicans(L.) Kuntze (38.29%) and was followed by Lonicera 
japonica Thunb, (37.14%) and Parthenocissus quinquefolia (L.) Planch. 12.57%). 
The most common nearest shrub species was Rubus argutus Link (25.71%) and 
was followed by Quercus alba L. (8.57%), Callicarpa americana L. (7.43%), and 
Quercus hemisphaerica Bartr. ex Willd. (6.86%). The most common nearest tree 
species with smaller dbh (5 inch or less) were Pinus taeda L. (30.86%) and Ilex 
vomitoria Ait. 12.00%) and with the larger dbh (> 5inch dbh) were Liquidam- 
bar styraciflua L. (23.43%), Triadica sebifera (L.) Small (20.00%), and Quercus 
alba L. 17.14%). 


ares 


ALLEN ET AL PLATANTHERA 


Taste 1. List of species associated with 35 clumps of green fringed orchid in Vernon Parish, Louisi- 
ana, during May 2005 with percent! of nearest individual records. 


ACANTHACEAE 
Ruelli liniensis (J.F.Gmel.) Steud. (1.14%) 


ACERACEAE 

Acer rubrum L. (5.71%, 1.71%, 
ANACARDIACEAE 

Toxi ron radicans (L.) Kuntze (38.29%) 


1.71%) 


APIACEAE 

Ptilimni apillaceum (Michx.) Raf. 
Sanicul densis L.(8.57%) 
AQUIFOLIACEAE 

llex opaca Ait. (1.14%, 0, 0) 

Ilex vornitoria Ait. (3.43%, 12.00%, 0) 
ARACEAE 

Arisaema triphyllum (L.) Schott 
ARALIACEAE 


Aralia spinosa L. (2.29%, 3.43%, 0.57%) 
ARISTOLOCHIACEAE 
Aristolochia serpentaria L. 
ASPLENIACEAE 
Asplenium platyneuron (L. 
ASTERACEAE 
Elephantopus carolinianus Raeusch. (2.86%) 
Helenium flexuosum Raf. 
/ t aAnnaden 


B.S.P. 


er 


lid anadensi L. 8 
So dace rugosa P.Mill. (1. 
Symphyotrichum eae (L.) A.& D. Love 
(2.29%) 

BETULACEAE 
Carpinus caroliniana Walt. (1.71%, 3.43%, 1.71%) 
Ostrya virginiana (P. Mill.) K. Koch (0, 1.14%, 0) 
a aan 

Big 1L. (3.43%) 
CAMPANULACEAE 
| icpilata 


heliaq annen 
ak 


CAPRIFOLIACEAE 

Lonicera japonica Thunb. (37.14%) 
Sambucus mye , cai: 2.29%, 0) 
Vib rife 


Vib dep ertune lt 


14%, 2.29%, 0) 


CELASTRACEAE 
Euonymus americana L. (0.57%, 0, 0) 


CLUSIACEAE 
Hypericum hyperi oldes (L.) Crantz 


Hynericum mutilum L 
P , 


CORNA 
Cornus pa [ L.(0.57%, 0, 0) 


CUPRESSACEAE 
Juniperus virginiana ee (0, 0.57%, 0) 


CYPERACEAE 

Carex complanata Torr. & Hook. (2.86%) 
Carex debilis Michx. 

Carex digitalis Willd. 

Carex festucacea Schkuhr ex Willd. 
Carex flaccosperma Dewey (0.57%) 
Carex longii Mack. (0.57%) 

Carex rosea Willd. (1.14%) 

ge ola ot: sh 


ea iao laantha ree 
DIOSCOREACEAE 
Dioscorea villosa L. 
DRYOPTERIDACEAE 
Onoclea sensibilis L. 
Palystichum acrostichoides (Michx.) Schott. 


EBENACEAE 
Diospyros virginiana L. (0.57%, 0, 0) 
ERICACEAE 

eee eee Marsh. 

tii Chapman (1. 


14%, 0, 0) 


EUPHORBIACEAE 
Triadica sebifera (L.) Small (1.14%, 8.00%, 20.00%) 


FABACEAE 

Galactia volubilis (L.) Britton 

Lespedeza cuneata (Dum.-Cours.) G.Don (0.57%) 

Trifolium repens L. (0.57%) 

Vicia sativa L.ssp. nigra (L.) Ehrh. (1.71%) 

FAGACEAE 

Fagus grandifolia Ehrh. (0, 0.57%, 10.29% 

Quercus alba L. (8.57%, 0.57%, 17.14%) 

Quercus hemisphaerica Bartr. ex Willd. (6.86%, 
6.86%, 0) 


—s 


Quercus nigra L. (4.00%, 2.29%, 1.14%) 


HAMAMELIDACEAE 
Hamamelis virginiana L. (0, 1.71%, 0) 


Tage 1.continued 


BRIT.ORG/SIDA 22(1) 


Liquidambar styraciflua L.(3.43%, 5.14%, 23.43%) 


JUGLANDACEAE 

Carya alba (L.) Nutt. ex Ell. (0, 0, 0.57%) 

Carya glabra (P. Mill.) Sweet var. hirsuta (Ashe) 
Ashe (5.14%, 0, 3.43%) 

Carya illinoinensis (Wangenh.) K. Koch 

JUNCACEAE 

alice coriaceus Mack. 

ia _ 57%) 


epha I) 


Lincs nol 


JUNCUS 16 uis Willd. : 7 ne 
LAMIACEAE 
Pycnanthemum tenuifolium Schrad. 
Salvia lyrata L.(0.57% 
Scutellaria elliptica Muhl. ex Spreng. (0.57%) 
LAURACEAE 
Sassafras albidum (Nutt.) Nees (0.57%, 0, 0) 
LILIACEAE 

inad ie a 00%) 
im michauxii 
Polygonatum bi oe (Walt.) Ell. 
LINACEAE 
Linum striatum Walt. 
LOGANIACEAE 
Gelsemium sempervirens (L. 


YCOPODIACEAE 


ma 


St. Hil. (0.57%) 


i 


| 
lygodium japonicum (Thunb. ex Murr.) Sw. 
(0.57%) 


MELIACEAE 

Melia azedarach L. (0.57%, 2.29%, 0.57%) 
MORACEAE 

Morus rubra L. (0.57%, 
MYRICACEAE 
Morella cerifera (L.) Small (0.57%, 0.57%, 0) 
NYSSACEAE, 

Nyssa sylvatica Marsh. (4.57%, 0, 4.00%) 
OLEACEAE 


een saat us L.(0,0,0.57%) 
Ir. (4.57%, 8.00%, 0) 


1.71%, Q) 


ONAGRACEAE 
liiduyinia ite ifaly L. 
Ludwigia palustris (L.) Ell. 


OROBANCHACEAE 
Epifagus virginiana (L) W. Bart. 


OSMUNDACEAE 
Osmijnda ci 


Osmunda regalis L. 
OXALIDACEAE 

Oxalis stricta L. 

PINACEAE 

Pinus taeda L. (0, 30.86%, 10.29%) 
POACEAE 
Andr dii Vitman (1.14% 


=< 


Briza minor L. 
hasmanthium laxum 

Chasmanthium sessi Worun (Poir.) Yates (17.71%) 

Dichanthelium acuminatum (Sw.) Gould & C.A 
Clark var. lindheimeri (Nash) Gould & CA. 
Clark 

Dichanthelium boscii (Poir.) Gould & C.A. Clark 
(4.00%) 


Dichanthelium commutatum (J.A. Schultes) 
Gould (4.57%) 
hanthelium dichot 7 (L.) Gould (12.57%) 

rum (Lam.) Gould (5.71%) 

Leersia virginica Willd. 

Lolium perenne L. (2.29%) 

Melica mutica Walt. 

is (L.) Beauv. (2.29% 

Paspalum u urvi ie Steud. 


Dich nth lium la if 


Oplist AEN IC Hirtell 


oe 


a (Michx.) Scribn. (0.57%) 


RHAMNACEAE 
Berchemia scandens (Hill) K. Koch (1.71%) 


edna 

Crataegus marshallii Eqglest. (0.57%, 0, 0) 
ae serotina Ehrh. (0.57%, 0, 0) 

Rubus argutus Link (25.71%, 0, 0) 

Rubus trivialis Michx. (3.43%, 0, 0) 
RUBIACEAE 

Galium pilosum Ait. 

oa ium tinctori um i. (4.00%) 


ichx. 
Mitchella repens L. . 86%) 
SMILACACEAE 
Smilax glauca Wall (2.29%) 


mila \ Walt. 


Smilax pu 


Smilax rotundifolia L. 


ALLEN ET Al 809 


Tas_e 1.continued 


Smilax smallii Morong VERBENACEAE 
Smilax tamnoides L. Callicarpa americana L. (7.43%, 2.29%, 0) 
le salaries VIOLACEAE 
hii (Desv.) Morton Viola sororia Willd. (1.14%) 
ULMACEAE VITACEAE 
Ulmus alata Michx. (1.14%, 2.29%, 2.86%) Parthenocissus quinquefolia (L.) Planch. (12.57%) 
Ulmus americana L. (0,0, 1.71%) Vitis rotundifolia Michx. (4.00%) 
URTICACEAE 


Boehmeria cylindrica (L.) Sw. 


' Percent of nearest individual records = number of nearest ae records divided by 175 and mul- 
tiplied by 100 for each peels Cae The categories aceous, woody vine, shrub (woody 
non-vines shorter than six ft t percent), ie ee non-vines taller than six feet 
and five inch or less dbh) eee percent), and trees (woody non-vines taller than six feet and 
larger than 5 inch dbh) (third percent). 


ACKNOWLEDGMENTS 


We would like to thank Larry Magrath and Charles J. Sheviak for reviewing our 
article. 


REFERENCES 


Correll, D.S. 1947. Additions to the orchids of Texas. Wrightia 1:166-181. 

Corrett, D.S. 1978. Native orchids of North America north of Mexico. Stanford University 
Press, Stanford, California. 

Fiora of NortH America EprtoriaL Committee. 2002. Flora of North America: Volume 26; 
Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford Univ. Press, New York. 

Liccio, J.and A.O. Licaio. 1999. Wild orchids of Texas. Univ. of Texas Press, Austin. 

MacRosents, D.T. 1977. Additions to the Louisiana flora. Sida 7:220-222. 

MacRoserts, M.H. and B.R. MacRoserts. 1998. A note on Platanthera lacera (Michx.) G. Don 
(Orchidaceae) in Louisiana. Phytologia 84:38-39. 

Priccon, A.M.and L.R. Ursatscu. 1977. Contributions to the flora of Louisiana. ||: Distribution 
and identification of Orchidaceae. Castanea 42:293-304. 

Ravroro, A.E., H.E. Antes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolina. 
Univ. North Carolina Press, Chapel Hill. 

Riovett, J.L. 1852. Catalogus Florae Ludovicianae. New Orleans Med. Surg. J. 8:734—754. 

THomas, R.D. and C.M. Auten. 1993. Atlas of the vascular flora of Louisiana, Vol. 1:ferns and 
ferns allies, conifers, and monocotyledons. Louisiana Department of Wildlife and Fish- 


eries, Baton Rouge. 

Soi Survey Starr. 2003. Soil survey of Vernon Parish, Louisiana. United States Department 
of Agriculture, Natural Resources Conservation Service, Washington, D.C 

USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National 
Plant Data Center, Baton Rouge, LA 70874-4490 USA. 


BRIT.ORG/SIDA 22(1) 


Book Notices 

Timber Press Books (Reviews Forthcoming) 

Davip Fross and Dirter WILKEN. 2006. Ceanothus. (ISBN 0-88192-762-7, 978-0- 
88192-762-7, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, 
Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, 
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). 
$39.95, 272 pp., color photos, b/w illustrations, 7" x 9". 


Contents.—Pretace, Acknowledgments. 1) Ceanothus in the Garden and Landscape (Natural Distri- 
bution, History of Cultivation, Growing Ceanothus 


Disease and Pests, Cultivar Development, Propa- 
gation, Ceanothus pecs ane Cultivars tox the Gi ina and L peeaes Ceanothus Selection Guide. 
2) i inthe I 
C ; 


stribution, Form and Function in Ceanothus. 


s, Other Ceanodus Names). C onversion Tables, Glossary, Bibliography, Index. 


aS 


Davin D. Stuart. 2006. Buddlejas. (ISBN 0-88192-688-4, 978-0-88192-688-0, hbk). 
Royal Horticultural Society Plant Collector Guide. Royal Horticultural Society 
and Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204- 
3527, US.A. (Orders: www.timberpress.com, mail@timberpress.com, 503- 
227-2878, 1-800-327-5680, 503-227-3070 fax.). $34.95, 192 pp., color pho- 
tos. a. 

Contents.—Foreword by Michael A. Dirr, Preface, Introduction. 1) Species. 2) Hybrids and Cultivars. 

3) Care and Cultivation. 4) Propagation. 5) Wildlife Attracted to Buddlejas. Where to See Buddlejas, 

Where to Buy Buddlejas, Bibliography, Index. 


RoBIN Wuite. 20060. Daphnes: A Practical Guide for Gardeners. (ISBN 0-88192- 
752-X, 978-0-88192-752-8, hbk.). Timber Press Inc, 133 S.W. Second Avenue. 
Suite +50, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimbet press.com 
mail@timberpress.com, 503-227-2878, l-800-327-5680, 503-227-3070 fax.). 
$34.95, 232 pp., color photos, 7" x 9". 


Contents.—Foreword, Preface, 1) History, Taxonomy and Mor = hology. 2) Daphne Species. 3) Daphne 
Hybrids. 4) Propagation. 5) Cultivation. 6) Pests and Diseases. 7) Garden Use. Useful Addresses. Glos- 
sary, Bibliography, Index of Daphnes, Conversion Tables. 


SIDA 22(1): 810. 2006 


SEYMERIA FALCATA (SCROPHULARIACEAE), 
A NEW RECORD FOR TEXAS AND THE UNITED STATES 
Joselyn Fenstermacher 
Sul Ross State University 


Alpine Texas 79832, U.S.A. 
josfenster@alumni.duke.edu 


ABSTRACT 


Upon review of herbari i | Hlecti itis clear that Seymeria falcata B.L. Turner 
irst report of 


occurs in the Dead Horse Mouncats of southern Brewster County, Texas. This is the f 
this species for Texas and the United States. 


RESUMEN 
Basandonos en la revision de especimenes herbario y colecciones recientes esta claro que Seymeria 
falcata B.L. Turner ocurre en las Montanas Caballos Muertos en el condado meridional de Brewster, 


Tejas. Este es ne primera cita de esta especie para Tejas y los Estados Unidos 


Turner (1982) provided a revisional study of the largely Mexican genus 
Seymeria. In this he described S. falcata B.L. Turner, a new species then known 
only from north-central Mexico, partially sympatric with its closest ally S. 
scabra Gray. The latter is typified by material collected by Charles Wright in 
the Davis Mountains of Jeff Davis County, Texas, and is distinguished by mostly 
symmetrical, semi-falcate fruits and markedly scabrous stems and foliage. In 
contrast, S. falcata has weakly to decidedly falcate fruits and minutely 
puberulous or glabrous stems and foliage, and glabrous fruits. 

Turner et al. (2003) recognized only a single species of Seymeria, S. scabra, 
as occurring in Trans-Pecos Texas. Recent collections from the Dead Horse 
Mountains in the Sierra Del Carmen of Big Bend National Park have revealed 
the presence there of S. falcata. 


Voucher specimens: TEXAS. Brewster Co.: Dead Horse Mountains, on rocky ridge above Brushy Can- 
yon, ca. 1/2 mi N of lodge, 1220 m, 15 Aug 1994, Talbot 137 (SRSC). Dead Horse Mountains, north 
sister of Sue Peaks at crest, 1740 m, 16 Sep 1994, Talbot 255 (SRSC); western slopes of Dead Horse 
Mountains below Sue Peaks, 1420 m, 3 Sep 2005, Fenstermacher 1187 (SRSC); Dead Horse Mountains, 
5 miinon Telephone Canyon Trail from Old Ore Road, 1310 m, 8 Mar 2005, Fenstermacher 571(SRSC). 
Seymeria falcata was first collected by Miller Talbot in the Dead Horse Moun- 
tains but his 1994 specimen was identified as Seymeria scabra. My recent col- 
lections of S. falcata from the Dead Horse Mountains spurred a review of the 
Seymeria specimens in the SRSC herbarium, thereby discovering and correctly 
identifying Talbot's previous collection. My S. falcata collections were made at 


SIDA 22(1): 811 — 812. 2006 


812 BRIT.ORG/SIDA 22(1) 


two locations in relative proximity to Talbot's collection sites, at elevations be- 
tween 1310 m and 1430 m (Fenstermacher 571, 1187, SRSC). | also collected S. 
scabra in a range of elevations in proximity to all collections mentioned here 
(Fenstermacher 744D, 1018, 1050, SRSC). The two species are clearly sympatric 
and hybridization between the two is suspected (Fenstermacher 1018, 1050, 
SRSC) 


ACKNOWLEDGMENTS 


| would like to acknowledge the generosity of Billie L. Turner and thank him 
for bringing this to my attention. Thanks also go to the Sul Ross State Univer- 
sity herbarium for research space, holdings, and technical expertise, without 
which this discovery would not have been made. The two reviewers, Guy L. 
Nesom and Richard D. Worthington, made helpful suggestions. 


REFERENCES 


Turner, B.L. 1982. Revisional treatment of the Mexican species of Seymeria 
(Scrophulariaceae). Phytologia 51:403-422. 

Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. 
Sida Bot. Misc. 24, Volume 1.Botanical Research Institute of Texas, Fort Worth. 


CAYRATIA JAPONICA (VITACEAE) NEW TO 
NORTH CAROLINA AND AN UPDATED KEY TO THE 
GENERA OF VITACEAE IN THE CAROLINAS 


Alexander Krings Robert J. Richardson 
Herbarium, Department of Botany Department of Crop Science 
North Carolina State University North Carolina State University 
Raleigh, North Carolina 27695-7612, U.S.A. Raleigh, North Carolina 27695-7620, U.S.A. 
Alexander_Krings@ncsu.edu Rob_Richard 


ABSTRACT 


A population of the introduced and expanding Cayratia japonica (Thunb.) Gagnep. (Vitaceae) is 


reported for North Carolina. Voucher specimens are cited and an updated key to the genera of Vitaceae 
in the Carolinas is presented. 
RESUMEN 


Se cita de Carolina del N blacién de | 1 


nexpansion Cayratid japonica 
(Thunb.) Gagnep. (Vitaceae). Se ae los pliegos tstigs y se presenta una clave puesta al dia de los 
géneros de Vitaceae en las Carolinas. 


The Asian vine Cayratia japonica (Thunb.) Gagnep. var. japonica (Vitaceae) is 
known from Louisiana, Mississippi, and Texas, but has not been previously re- 
ported escaped elsewhere in the United States (Shinners 1964; Brown 1992; Al- 
ford 2003; USDA, NRCS 2005). The species was not previously reported for North 
Carolina by Radford et al. (1968), Pittillo et al. (1969), Pittillo et al. 1972), Kral 
(1981), Pittillo and Brown (1988), or USDA, NRCS (2005). However, an aggres- 
sive and persistent population of the plant was recently encountered on pri- 
vate property in a suburban area near Winston-Salem (Forsyth Co., NC). Vines 
were growing up trees (reaching heights of 9-12 m high), along a fence, and 
generally throughout the landscape—covering about 0.5 acre. When visited in 
late August, the vines were in full flower and well-attended by honeybees and 
wasps. Based on conversations with the landscaper, individual ramets aggres- 
sively re-sprouted following handweeding. Apparently, a vine of > 2.5cm diam 
at the base had been removed. Remaining roots at the site were ca. 0.6 cm diam. 
It remains unclear how the species arrived. Both previous and current 
homeowners, the latter an avid birdwatcher, indicated that the species had not 
been planted. However, intentional planting or contamination through other 
ornamental plantings cannot be ruled out. Bird dispersal may be a possibility, 
although unlikely as additional populations between North Carolina and the 
Gulf Coast would be expected under this scenario and reproduction is reported 
to be primarily vegetative in North America—at least in Texas (Brown 1992). 


SIDA 22(1): 813 - 815. 2006 


BRIT.ORG/SIDA 22(1) 


Although the vector of arrival remains unknown, the persistence of the North 
Carolina population is of concern. Alford (2003) reported a Mississippi popu- 
lation tripling in size within LO months. Little information is currently avail- 
able regarding control measures. 

Voucher specimens: U.S.A. NORTH CAROLINA. Forsyth Co.: Wesleyan Lane, Winston-Salem, 1] Aug 


2005, Jonathan Todd s.n.ONCSC, USCH). Wesleyan Lane, Winston-Salem, 29 Aug 2005, Robert]. Rich- 
ardson s.n. (FLAS, NCSC, USCH). 


Cayratia japonica (Fig. 1) can be distinguished from other Vitaceae taxa in the 
Carolinas by the pedate quinqueloliate leaves. Tendrils are bifurcating and not 
disk-tipped. An updated key to the genera of Carolina Vitaceae follows. Cissus 
is included here based on the listing of C. trifoliata (L.) L. by Weakley (2005). 
Cissus trifoliata is cultivated in North Carolina, but-has not been observed to 
escape (Krings, pers. obs.). 
1. Tendrils, at least some, terminating in adhesive disks 
1. Tendrils lacking adhesive disks 
2. Leaves pedate quinquefoliate 
2. Leaves simple, palmately, or pinnately compound, but never pedate 
quinquefoliate. 
3. Inflorescence a thyrse; petals connate at their apices Vitis L. 
3. Inflorescence a dichotomous or umbelliform cyme; petals free at their apices. 
4. Cyme dichotomous; flowers 5-merous 
4. Cyme umbelliform; flowers 4-merous 


Parthenocissus Planch. 


Cayratia Juss. 


OQ. 


Ampelopsis Michx. 
Cissus L. 


ACKNOWLEDGMENTS 


We thank Jonathan Todd for bringing the Forsyth population to our attention 
and the curators and staff of DUKE and NCU for access to their collections. 
Guy Nesom offered helpful review comments. 


REFERENCES 


Atrorb, M.H. 2003. Noteworthy collections Mississippi Castanea 68:93. 


Brown, L.E. 1992. Cayratia japonica (Vitaceae) and Paederia foetida (Rubiaceae) adventive 
in Texas. Phytologia 72(1): 45-47. 


Kral, R.1981.Some distributional reports of weedy or naturalized foreign species of vas- 
cular plants for the southern states, particularly Alabama and middle Tennessee. Cas- 
tanea 46:334-339. 

Pittito, J.D., JH. Horton, and K.W. Greentee. 1969. Additions to the vascular flora of the Caro- 
linas. |. J. Elisha Mitchell Sci. Soc. 85:18-22. 

Prrmitto, J.D., JH. Horton, and K.E. Herman. 1972. Additions to the vascular flora of the Caroli- 
nas. ll. J. Elisha Mitchell Sci. Soc. 88:144-152 

Pirtito, J.D.and A.E. Brown. 1988. Additions to the vascular flora of the Carolinas. Ill.J. Elisha 
Mitchell Sci. Soc. 104:1-18. 


Raprorod, A.E., H.E. AHves, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
The University of North Carolina Press, Chapel Hill. 


KRINGS AND RICHARDSON, CAYRATIA JAPONICA NEW TO NORTH CAROLINA 


foli leaf 1 bif Jril: B. inflorescence: C. detail 
Hy ec 


14:-L t 1 dol} 


/ | 1) ond €] hid-M dick (ctl 1). Leaf, flower bud, 


on? (NCU); 4ril 1 inf] I d S fell 1580 (NCU). 


SHinners, L.H. 1964. Cayratia japonica (Vitaceae) in southeastern Louisiana: new to the 
United States. Sida 1:384. 

USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). Data com- 
piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge. 

Weaxtey, A.S.2005.Flora of the Carolinas, Virginia,and Georgia:Working draft, 10 June 2005. 
North Carolina Botanical Garden, Chapel Hill. 


BRIT.ORG/SIDA 22(1) 


Book NOTICES 


Blackwell Publishing 
Kenpaut R. LaMkKey and Micnaet Ler. (Eds.). 2006. Plant Breeding: The Arnel R. 
Hallauer International Symposium. (ISBN 0-8138-2824-4, 978-0-8138-2824- 
4, hbk.). Black well Publishing, 2121 State Ave., Ames, [A 50014-8300, U.S.A. 
and 9600 Garsington Road, Oxford, OX4 2DQ, UK. (Orders: 515-292-0140, 
515-292-3348 tax 1-800- oe 6657, www.blackwellprofessional.com, 
lers@ blackwell hing.com). $149.99, 379 pp., numerous illus- 


trations, 7" < LO", 

Douc-as D. Stokke and Lestir H. Groom. 2006. Characterization of the Cellulosic 
Cell Wall. (ISBN 0-8138-04 39-6, 978-0-81 38-04 39-2. hbk.). Black well Pub- 
lishing. 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington 
Road, Oxford, OX4 2DQ, UK. ue 515- fatal 0140, 515-292-3348 fax l- 


800-862-6657, www.black well ional.com, ord blackwell- 
publishing.com). $149.99, 274 Dp, color photos, numerous b/w illustrations, 
7" X 10" 


JouN F Lesuir and Brett A. SUMMERELL. 2006. The Fusarium Laboratory Manual. 
(ISBN 0-8138-1919-9, 978-0-8138-1919-8, spiral pbk.). Blackwell Publishing. 
2121 State Ave., Ames, IA 50014-8300, U.S.A.and 9600 Garsington Road, Oxford, 
OAT 2ZDO.U - LOnderss ee 292-0140, 515-292-3348 fax 1-800-862-6657, 
blacl ional.com, orders@ blackwellpublishing.com). 


VV VV 
$124.99, 388 * b/w Tiveeetione Ole 
From the publisher—For the first time in over 20 years, a comprehensive collection of photographs 
and descriptions of species in the fungal genus Fusarium is available. This laboratory manual pro- 
vides an overview of the biology of Fusarium and the techniques involved in the isolation, identi 
cation and characterization of individual species and the populations in which they occur. It is the 
ogical and molecular approaches have been incorporated into a vol 


first time that genetic, morphol 
h new and 


ume devoted to Fusarium identification. The authors include descriptions of species, bot 
old, and provide protocols for genetic, morphological and molecular identification techniques. 


SIDA 22(1): 816. 2006 


NEW AND NOTEWORTHY COLLECTIONS FOR ARKANSAS 


James H. Peck Brett E.Serviss 
Department of Biology Department of Biology 
University of Arkansas at Little Rock Henderson State University 
Little Rock, Arkansas 72204, U.S.A. bags ia, Arkansas 71999, U.S.A. 
jhpeck@ualredu visb@hsu.edu 
ABSTRACT 
Ten species of vascular plants, all native or naturalized in the soutaeastern United States, are re- 
ported as new or noteworthy for Arkansas. Of these, fi -native. Seven are 
reported as state records, and the accounts for the other three species provide clarifications, includ- 


ing updates on the status of two aquatic weeds, one federally listed as an invasive non-native weed 

(Hydrilla verticillata) and another erroneously thought to be native to the United States (Salvinia 

ma). Species reported as a state record are: Amaranthus blitum, Chamaesyce hypercifolia, 

Chamaesyce opthalmica, Euphorbia graminea, Hedyotis corymbosa, Ipomoea quamoclit, and 
4 {] | 


FAFriLClare AYLUT LUM 
RESUMEN 
Se citan diez especies de plantas vasculares lizad te de los Estados 
dos bl Ark 


Uni i rkansas. De estas, Ci i y cl i Siete se 
acts : snes ] lds 


citan para el estado, y par 
oe estatus de dos ee acuaticas, una ade ™ as del listado federal de plantas 1 invasoras no nativas 


minima). Las especies citadas como nuevas ve ara el estado son: Amaranthus blitum, Chamaesyce 
hypercifolia, Cl : 1, Hedyotis corymbosa, Ipomoea quamoclit, 


[ fo) 


y Panevan oridana. 
INTRODUCTION 


The Arkansas Vascular Flora Project is an on-going collective effort to produce 
a complete and modern treatment to the vascular flora of Arkansas in the form 
of achecklist, atlas, and manual (Peck 2003). Toward that end, urban and rural 
parks, transportation routes, and other ruderal areas in Pulaski Co. and else- 
where in central Arkansas were surveyed to test the completeness of the floris- 
tic record. The state occurrence and state county distribution of species col- 
lected was determined from dot-maps kept at Herbarium LRU, University of 
Arkansas at Little Rock. The national occurrence and distribution of the new 
state records was determined from the national flora database kept by NRCS 
(1999). Records at the state and county level were documented with herbarium 
specimens. The records reported here support the notion that lesser quality 
habitats, such as urban and rural disturbed or ruderal habitats, whether xeric 
ones along railroad tracks or aquatic ones such as mudflats along impound- 
ments, are under-collected. Although less pristine and aesthetic than non-an- 


SIDA 22(1): 817 — 820. 2006 


818 BRIT.ORG/SIDA 22(1) 


thropogenic habitats, ruderal environments warrant further survey in Arkan- 
sas to ensure completeness of the floristic record. 

SPECIES NEW OR NOTEWORTHY FOR ARKANSAS 
Amaranthus blitum L. (Amaranthaceae). Purple amaranth, a US non-native, 
occurs across the coastal states from Massachusetts south to Florida and west 
to Louisiana and Texas. This is the first documentation of this species in Ar- 
kansas. 


Voucher specimen: Pulaski Co.: urban ruderal habitat, along railroad tracks at University Ave. and 
65" St, Little Rock, 13 Oct 2001, Peck 2001231 (LRU). 


Chamaesyce hirta (L.) Millsp. (Euphorbiaceae). Pillpod sandmat occurs across 
the coastal states from South Carolina south to Florida and west to Louisiana 
and Texas. Known from Bradley and Pope Cos., Arkansas. This is the third docu- 
mentation of this species in Arkansas. 

Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. and 
65" St. Little Rock, Ll Nov 2001, Peck 20014892, 2001499, 20014895, 2001503 (LRU). 


Chamaesyce hypercifolia (L.) Millsp. Euphorbiaceae). Graceful sandmat occurs 
across the southeastern United States west to Louisiana and Texas. This is the 
first documentation of this species in Arkansas. 


Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. anc 
65" St, Little Rock, 10 Nov 2001, Peck 2001490, 2001496, 2001500, 2001504 (LRU). 

Chamaesyce opthalmica (Pers.) Burch (Euphorbiaceae). Florida hammock 
sandmat occurs in Georgia and Florida west to Louisiana. This is the first docu- 
mentation of this species in Arkansas. 

Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. and 
65" St. Little Rock, 10 Nov 2001, Peck 2001488, 2001497, 2001502 (LRU 

Euphorbia graminea Jacq. (Euphorbiaceae). Grassleal spurge, a US non-native, 
occurs as escapes in Florida and Hawaii. This is the first documentation of this 
species in Arkansas. 


— 


Voucher specimens: Pulaski Co.: urban ruderal habitat, along railroad tracks at Barton state fair 
grounds, Roosevelt Rd., Little Rock, 1] Nov 2001, Peck 2001567 (LRU): ruderal Saita near railroad 
tracks, east of University Ave. at 65" St, Little Rock, 4 Sep 1999, Peck 99764 (LRL 

Hedyotis corymbosa (L.) Lam. [Oldenlandia corymbosa L_] eee: Flat- 
top mille graines is a coastal species from South Carolina south to Florida and 
west to Louisiana and Texas. This is the first documentation of this species in 
Arkansas. 


Voucher specimens: Jefferson Co.: urban ruderal habitat, along railroad tracks at Old 167 in Pine 
Bluff near Arkansas River, 14 Oct 2001, Peck 2001243. Pulaski Co.: ruderal habitat, along railroad 
tracks at University Ave. and 65" St., Little Rock, 13 Oct 2001, Peck 2001232 (LRU). Saline Co.: ruderal 
ditat, along railroad tracks, S side of Benton, 13 Oct 2001, Peck 2001236 (LRU). 


— 


ha 


PECK AND SERVISS, NOTEWORTHY COLLECTIONS FOR ARKANSAS 819 


Hydrilla verticillata (L. f.) Royle (Hydrocharitaceae). Waterthyme, United States 
non-native and an aggressive invasive species in the southeastern United States, 
especially from Florida west to Texas, was first documented from Arkansas by 
Parker and Serviss (2003). The present collections further document the spread 
of this species downstream along the Ouachita River impoundments to other 
counties in central Arkansas. 


Voucher specimens: Garland Co.: stranded on mudflat by drawdown event of Ouachita River, shore- 
line at boat ramp park at N side of Carpenter Dam, Lake Hamilton, 23 Nov 2004, Peck 2004248 (LRU); 
stranded on mudflat by drawdown event of Ouachita River, shoreline at Spencer’s Bay, mouth of 
Gulpha Creek, Lake Catherine, 23 Nov 2004, Peck 2004240 (LRU); stranded on mudf a by draw- 
down event on Ouachita River, Lake Ouachita, shoreline at Brady Mountain campground, end of CR 
119, 9 Dec 2004, Davis s.n. (LRU). Hot Spring Co.: stranded on mudflat by Eerie n event of Oua- 
chita River, shoreline of Lake Catherine, Lake Catherine State Park, 23 Nov 2004, Peck 2004243 (LRU). 
Montgomery Co.: stranded on mudflat by drawdown event of Ouachita River, shoreline at Big Fir 
Campground, Lake Ouachita, end of CR 56, 23 Nov 2004, Peck 2004262 (LRU). 


Ipomoea quamoclit L. (Convolvulaceae). Cypressvine is native to tropical 


America, but is known to escape from cultivation across the southeastern 
United States, and is known from all states surrounding Arkansas. This collec- 
tion documents the first occurrence of this species in Arkansas. 


Voucher specimen: Pulaski Co.: urban ruderal habitat, present for three years as escape at informal 

ae grounds along railroad tracks at Barton state fair grounds, Roosevelt Rd., Little Rock, 14 
Nov 2005, Peck 2005009 (LRU). 

Parietaria floridana Nutt. (Urticaceae). Florida pellitory is a coastal species that 

occurs from Delaware south to Florida and west to Louisiana and Texas. This is 

the first documentation of this species in Arkansas. 


Voucher specimens: Garland Co.: ruderal habitat along roadside, 4 mi E Hot Springs on 270, 22 May 
2002, Peck 2002043 (LRU). Grant Co.: ruderal habitat along roadside, 3 mi E Poyen, 22 May 2002, 

Peck 2002032 (LRU). Hot Spring Co.: ruderal habitat along roadside, 4 mi S Malvern on 9, 22 May 
2002, Peck 2002043 (LRU). Pulaski Co.: urban ruderal habitat along roadside in Boyle Park, Little 
Rock, 22 May 2002, Pec : 2002018 (LRU). Saline Co.: ruderal habitat along roadside, 4 mi S Benton on 
25, 22 May 2002, Peck 2002054 (LRU). 


Salvinia minima Baker (Salvinieaceae). Least waterspangles, a non-native US 
aquatic fern, occurs in the southeastern United States from South Carolina south 
to Florida and west to Louisiana and Texas (Peck, 2002). The species was first 
documented from Arkansas by Peck (1999, 2002). These recent collections docu- 
ment a westerly range extension to three counties in west-central Arkansas. 

Voucher specimens: Garland Co.: stranded on mudflat by drawdown event of Ouachita River, shore- 
line at boat ramp park at N side of Carpenter Dam, Lake Hamilton, 23 Nov 2004, Peck 2004250 (LRU); 
stranded on mudflat by drawdown event of Ouachita River, shoreline at Spencer's Bay, mouth of 
Gulpha Creek, Lake Catherine, 23 Nov 2004, Peck 2004242 (LRU); stranded on mudflat by draw- 
down event on Ouachita River, Lake Ouachita, shoreline at Brady Mountain Campground, end of CR 
119, 23 Nov 2004, Peck 2004260 (LRU). Hot Spring Co.: stranded on mudflat by drawdown event of 
Ouachita River, shoreline of Lake Catherine, Lake Catherine State Park, 23 Nov 2004, Peck 2004245 


820 BRIT.ORG/SIDA 22 


_~ 


1) 


(LRU). Montgomery Co.: stranded on mudflat by drawdown event of Ouachita River, shoreline at Big 
Fir Campground, Lake Ouachita, end of CR 56, 23 Nov 2004, Peck 2004264 (LRU). 


ACKNOWLEDGMENTS 


The authors thank Theo Witsell and one anonymous reviewer for making im- 
provements to the manuscript. 
REFERENCES 

NRCS, Nationat Resources CONseRVATION Service. 1999. The PLANTS database. [http:// 
plants.usda.gov/plants]. USDA, National Plant Data Center, Baton Rouge, LA . 

Parker, M. and B.E. Seaviss. 2003. Occurrence and status of Hydrilla verticillata (L. f.) Royle 

Hydrocharitaceae) in Arkansas. J. Arkansas Acad. Sci. 57:15. 

Peck, J.H. 1999. Salvinia minima in Arkansas. Amer. Fern. J.89:215-216. 

Peck, JH. 2002. Survey of Salvinia (Salviniaceae) in eastern Arkansas. J. Arkansas Acad. Sci. 
55:115-118. 

Peck, J.H.2003. Arkansas fl iditi instat ts,exclusions, and re-exclusions. Sida 
20:1737-1757. 


— 


CLEMATIS MOREFIELDII (RANUNCULACEAE) 


NEW TO TENNESSEE 
Dwayne Estes Chris Fleming 
The Herbarium Breedlove, Dennis, Young suean Inc. 
Dept. of Ecology & i cee Biology 133 Holiday Court / Suite 2 
The University of Tennessee Franklin, Tennessee 3706/7, 7 A. 
Knoxville, Tennessee 3 USA cfleming@bda-inc.com 


tnplants@yahoo.com 


ABSTRACT 


Clematis Cue (Morefield’s leather flower, Huntsville vasevine), a ieedetaly endanseree ape 


Ta 


nted for the flora of Tennessee for the first time 


a small ae ravine ee on the lower slopes of the western seionied of the Guabatiad 
Plateau in Franklin County, Tennessee. A general description of the habitat and associated flora is 
included. 


RESUMEN 


Clematis morefieldii (flor de cuero de Moretield,), especie en peligro, se documenta para la !lora de 


Sarasa BOE Paine. vez. Se descubri ieron dieciocho plantas a lo largo de un pequeno barranco en 
lental del Cumberland Plateau en Franklin County, Tennessee. 


Se incluye una descripcion pencil del habitat y de la flora asociada. 
INTRODUCTION 


Clematis morefieldii Kral (Morefield’s leather flower, Huntsville vasevine) is a 
federally endangered species previously thought to be endemic to Madison and 
Jackson counties, Alabama (Kral 1987; USFWS 1992; Pringle 1997). This rela- 
tively recently described species (Kral 1987) is similar to the widespread C. 
viorna but differs from all variants of that species in the cobwebby tomentose 
to villous pubescence of its stems, and in having bracts at or very near the base 
of the peduncle rather than well above the base (Pringle 1997). Clematis 
morefieldii also usually differs from C. viorna in having more flowers clustered 
in the leaf axils on shorter peduncles, in its sepals more pinkish and greenish- 
tinged, and abaxial leaf surfaces more densely pubescent (Kral 1987). 

As of 2003, C. morefieldii had been documented from ca. 10 populations, 
and currently at least two of these (including the type locality) are considered 
extirpated. All known C. morefieldii sites occur along the dissected western es- 
carpment of the Cumberland Plateau in rocky limestone woods on mostly south 
or southwest-facing slopes. These sites are typically dominated by Juniperus 
virginiana L. and various hardwood species characteristic of basic soils, includ- 
ing Cotinus obovatus Raf., which isa key indicator species for C. morefieldii (Kral 


SIDA 22(1): 821 — 824. 2006 


822 BRIT.ORG/SIDA 22(1) 


1987). This habitat type, referred to by Kral (1987) as the Cotinus association, is 
restricted to the lower slopes of the western escarpment of the Cumberland 
Plateau in northeastern Alabama and southeastern middle Tennessee. Kral 
(1987) suggested that further field exploration of suitable habitats within this 
community type in Alabama and Tennessee could result in the discovery of 
additional populations. 

In spring 2003, we began searching lor C. morefieldiiin southeastern middle 
Tennessee close to the nearest known Alabama population of C. morefieldii.On 
5 Jun 2003, we discovered a small population of C. morefieldii in Franklin 
County, Tennessee ca. 12.5 km northeast of the nearest Alabama population. 
This is the first report lor C. morefieldii from Tennessee. 

Voucher specimen: Franklin Co.: ca. 4.2 mi SE of Huntland (Beans Creek Quad), SE of Motlow Cove 
in headwaters of Wilhite Creek, N side of Blooin Hollow, cela Fee small seasonal stream in 
rocky limestone woods, 5 Jun 2003, D. Estes & C. Fleming 04795 N). 

DISCUSSION 
The Franklin County, Tennessee site is on private property approximately 6 
km ESE from the town of Huntland on the dissected western escarpment of 
the Cumberland Plateau in the Elk River watershed of the Tennessee River Ba- 
sin. The population is located along the lower portion of a SSW-facing slope 
over Monteagle limestone at an elevation of ca. 347 m (1140 ft). Eighteen plants, 
one in flower and one with immature fruit, were found. Most of the vines were 
0.5 m or less in length and apparently damaged by insect herbivory. The indi- 
viduals were scattered along a 20 m reach of a wet weather conveyance 1-2 m 
in width, growing both along the banks and among the boulder substrate within 
the channel. The surrounding forest contained the following associates: Acer 
saccharum Marsh., Carya carolinae-septentrionalis (Ashe) Engl. & Graebn., 
Celtis tenuifolia Nutt, Fraxinus americana L., F quadrangulata Michx., Quercus 
muehlenbergii Engelm., Juniperus virginiana L., Cotinus obovatus, Cercis 
canadensis |. Ostrya virginiana (P. Mill.) K.Koch, Forestiera ligustrina (Michx.) 
Poir, Hamamelis virginiana L., Hypericum frondosum Michx., Rhus aromatica 
Ait., Bumelia lycioides (L.) Pers., Symphoricarpos orbiculatus Moench, and the 
rare Neviusia dlabamensis Gray and Viburnum bracteatum Rehd. The most 
common herbaceous taxa observed were Dasistoma macrophylla (Nutt.) Raf. 
Polymnia canadensis L., Scutellaria ovata Hill, and Solidago auriculata Shuttlw 
ex Blake. 

Several species considered rare in Tennessee were discovered growing with 
or adjacent to the Clematis population, including a small population of the state- 
endangered (TNHP 2004) Viburnum bracteatum (Estes & Fleming 04788 
TENN). One shrub was growing at the edge of the wet weather conveyance con- 
taining C. morefieldii and another group of 15-20 individuals were ca. 100 m 


TO TENNESSEE 823 


away along the rocky banks of a nearby stream. This species is extremely rare 
throughout its range and is endemic toa small area centered on the Cumberland 
Plateau of northeastern Alabama, northwestern Georgia, and southeastern 
Middle Tennessee. This is the second reported Tennessee occurrence for V. 
bracteatum(C. Bailey, Tennessee Natural Heritage Program, pers. comm.). Out- 
side of Tennessee, the species is only known from ca. eight extant populations. 
There is one each in Floyd and Walker counties in Georgia J. Allison, Georgia 
Natural Heritage Program, pers. comm.), and six documented populations in 
Alabama (Etowah, Jackson, and Madison counties), although one of these has 
not been observed in more than 50 years (M. Barbour, Alabama Natural Heri- 
tage Program, pers. comm.). 

Another significant find at the site was a small population of the state- 
threatened (TNHP 2004) Neviusia alabamensis (Estes & Fleming04790 TENN). 
Approximately 30-40 stems were located along a rocky streambank in asso- 
ciation with C. morefieldii. The Neviusia, like the Clematis, was visibly impacted 
by herbivory. Our collection represents a new county record for N. alabamensis 
in the state and the first report for the species from the Tennessee portion of the 
Cumberland Plateau; however, other populations are known fror the 
Cumberland Plateau of Alabama (Long 1989; D. Estes, pers. obs.) and Georgia 
(Long 1989). There are nine previously reported Tennessee populations (C. Bailey, 
Tennessee Natural Heritage Program, pers. comm.) within the Central Basin 
Section or at the transition zone between the Central Basin and Highland Rim 
Sections of the Interior Low Plateaus (ILP) Physiographic Province (Horn & 
Somers 1981; Long 1989). The Franklin County station is ca. 18 km southeast of 
the nearest previously known Tennessee population in Moore County. 

Other rare taxa tracked by the Tennessee Natural Heritage Program (2004) 
found within 100 m of the C. morefieldii site were Juglans cinerea L. Tennessee 
threatened), Lonicera dioica L. Tennessee threatened), Solidago auriculata (Ten- 
nessee threatened), and Cotinus obovatus (Tennessee special concern). The 
Cotinus and Solidago were common components of the forest while only one 
individual was found of J. cinerea and L. dioica, respectively. 


SUMMARY 


The recent discovery of C. morefieldii in Tennessee not only represents a state 
record but also extends the range of the species by some 12.5 km to the north- 
east. Recent fieldwork in the area has revealed several sites with suitable habi- 
tat for C. morefieldii on the western escarpment of Tennessee’s Cumberland Pla- 
teau from the Alabama state line northeastward to southern Warren County. 
Systematic surveying in this area might result in the discovery of more popula- 
tions of the federally endangered Morefield’s leather flower as well asa number 
of other significant rare taxa. 


824 BRIT.ORG/SIDA 22(1) 


ACKNOWLEDGMENTS 


Our sincere gratitude is extended to Drs. Robert Kral and Michael Dennis for 
confirming our identification of C. morefieldii. Furthermore, we appreciate the 
information provided to us by Steve Threlkeld of the Alabama Department of 
Conservation and Natural Resources, Al Schotz and Michael Barbour of the 
Alabama Natural Heritage Program, James Allison of the Georgia Natural Heri- 
tage — and Claude Bailey of the Tennessee Division of Natural Heritage. 
Thanks are also due to Dan Spaulding, curator of collections at the Anniston 
Museum of Natural History, who loaned us specimens of Morefield’s leather 

flower from the nearest Alabama population. Dr. B. Eugene Wofford, curator of 
the University of Tennessee Herbarium, was very helpful throughout the project 
and his editorial comments improved the manuscript. We wish to thank the 
Department of Botany at the University of Tennessee for supporting the travel 
costs associated with this project. 


REFERENCES 


SS 


Horn, D.D.and P. Somers. 1981. Neviusia alabamensis (Rosaceae) in Tennessee. Sida 9:90-91, 

Krat, R. 1987. A new “Viorna” Clematis from northern Alabama. Ann. Missouri Bot. Gard. 
74:665-669, 

Lona, A.A. 1989. Disjunct populations of the rare shrub, Neviusia alabamensis Gray (Ro- 
saceae). Castanea 54:29-39, 

Prinate, J. 1997. In: Flora of North America Editorial Committee, eds. Flora of Nort 
North of Mexico. Vol. 3. Oxford University Press, New York. 

TENNESSEE NATURAL HERITAGE PROGRAM (TNHP). 2004. Tennessee rare plant list. Tennessee Divi- 
sion of Natural Heritage, Dept. Environ. Conserv., Nashville. 

United States Fish and Witouire Service (USFWS). 1992. Endangered and threatened wildlife 
and plants; determination of Clematis morefieldii (Morefield’s leather flower) to be an 

endangered species. Fed. Register 57(98):21562-34420. 


1 America 


GRATIOLA BREVIFOLIA (PLANTAGINACEAE) NEW TO THE 
FLORA OF DELAWARE, THE DELMARVA PENINSULA, 
AND THE MID-ATLANTIC 


Wesley M. Knapp Dwayne Estes 
Maryl ag aie ‘ment of Natural Resources Univers! ee Mee 
Idlife & Heritage Service a utionary Biology 
PO Box 68 xville Se 37996, USA. 
Wye Mills, Maryland 21679, U.S.A. tnplants@yahoo.com 


wknapp@dnr.state.md.us 


ABSTRACT 
Gratiola brevifolia (Pl i d | native additi he fl f Delaware, 
the Delmarva Peninsula, arid tarhe Mid- Atlantic This species is dis} PI ly 835 km (520 


mi) from the closest known population in Burke Co., Georgia. 


RESUMEN 


Gratiola brevifolia (Plantagina 


e cita como una gone native rara ea anadir a la flora de 
835 km (520 


Delaware, la Peninsula de Delmarva, y peuane Minds Atla 
mi) aproximadamente de la poblaci cida cana €1 Burke Co; Georgia, 


The genus Gratiola L. (Plantaginaceae) consists of ca. 30 species (Estes, unpub- 
lished data) widely distributed through the North and South Temperate zones 
and on mountains within the Tropics (Pennell 1935). Six species of Gratiola 
(s.) have been reported on the Delmarva Peninsula (Tatnall 1946, McAvoy 2001), 
an area that lies entirely within the Atlantic Coastal Plain Physiographic Prov- 
ince of the eastern United States and consists of the Eastern Shores of Mary- 
land and Virginia and the majority of the state of Delaware. The northern por- 
tion of Delaware lies within the Piedmont Physiographic Province (Plank & 
Schenck 1998). These six species are G. aurea Muh. G. neglecta Torr, G. pilosa 
Michx., G. ramosa Walt., G. virginiana L.,and G. viscidula Pennell. On Delmarva, 
G. pilosa and G. virginiana are considered to be common, G. aurea and G. 
neglecta infrequent, and G. ramosaand G. viscidula are thought to be historical, 
having not been reported for 20 or more years (McAvoy, pers. comm.). Gratiola 
ramosa was last collected from Wicomico Co., Maryland, near the town of Sal- 
isbury (Canby s.n. PH), and G. viscidula was last collected from New Castle Co., 
Delaware, near the city of Wilmington (Tatnall s.n. DOV; Commons s.n. DOV). 
On 23 June 1992, Frank Hirst and Ron Wilson reportedly rediscovered 
Gratiola ramosa in Sussex Co, Delaware (R. Wilson 0719941, pers. herbarium). 
Recently, this population was visited by the first author and specimens were 
collected and sent to the second author for verification. Subsequently, the speci- 


SIDA 21): ): 825 — 829. 2006 


826 BRIT.ORG/SIDA 22(1) 


mens were determined not to be G. ramosa but instead were identified as G. 
brevifolia Ral., a species not previously reported from Delaware, the Delmarva 
Peninsula, or the Mid-Atlantic. Though G. brevifolia and G. ramosa are easily 
distinguished by the presence or absence of two bracteoles at the base of the 
calyx (Godt rey @ Wooten 1981), Hirst and Wilson’s misidentilication could be 
expected because G. brevifolia is not included in any manual commonly used 
in the Mid-Atlantic Ge., Fernald 1950; Radford et. al 1968; Brown & Brown 1984: 
Gleason & Cronquist 1991). 
Voucher Specimen: U.S.A, DELAWARE. Sussex Co.: Coastal Plain Physiographic Province, ditched 
portions of Tussocky Branch paralleling Piney Branch Rd., northwest of the town of Delmar, 29 Jul 
2005, Knapp 1549 (DOV, TENN, Maryland Natural Heritage Program Herbarium). 


Gratiola brevifolid is an erect perennial of wet, sandy pinelands (Pennell 1935), 
oak barrens (Tennessee), and sandy riverbanks (Arkansas and Oklahoma). The 
species ranges from Tennessee and Florida west to Oklahoma and Texas (Pennell 
1935) and is found in four centers of distribution, with two occurring east and 
west of the Mississippi (Fig. L). Eastward, G. brevifolia is centered on the Coastal 
Plain of northern Florida and southern Georgia with scattered populations west 
tosoutheastern Alabama. The species is also significantly disjunct to the High- 
land Rim and Cumberland Plateau of central Tennessee (Chester et al. 1997). To 
the west, G. brevifolia is most frequent in the West Gulf Coastal Plain of south- 
eastern Texas and southwestern Louisiana, with disjunct populations in the 
West Gulf Coastal Plain of southeastern Oklahoma and the Ouachita Province 
of Oklahoma and Arkansas (Estes, unpublished data). Despite the broad geo- 
graphic distribution, it is considered imperiled in Arkansas (NatureServe Ex- 
plorer 2005) and probably should be considered a species of conservation con- 
cern in Alabama, Oklahoma, and Tennessee (Estes, unpublished data). 

ld ramosdasa species of conservation concern 


pa: 


for Delaware based upon the discovery and identification by Hirst and Wilson. 
With this publication, G. ramosd should no longer be considered a component 
of Delaware's native flora; however, it was historically a component of 
Delmarva’s native flora based on Canby’s collection cited above. 

The population of Gratiola brevifolia in Delaware consists of ca. 1500 in- 
dividuals restricted to the lower banks and bottoms of a ditched perennial 
stream. Interestingly, several other species in the vicinity of the G. brevifolia 
population are considered rare in Delaware, based on McAvoy (2003). These 
species include: Amphicarpum purshii Kunth, Hypericum adpressum Raf. ex 
WeBart., H. denticulatum Walt., Sabatia difformis (L. Druce, and Utricularia 
rddiata Small. 

The discovery of Gratiola brevifolia in Delaware marks a new addition to 
the flora of Delaware, the Delmarva Peninsula, and the Mid-Atlantic region, 
and is significantly disjunct from all other known populations in the South- 
east. The closest population to the Delaware occurrence is ca. 835 km (520 mi) 


KNAPP AND ESTES PNR WHERES 997 


Fic. 1. Range of Gratiola brevifolia. Each @ represents a specimen examined. 


to the southwest in Burke Co., Georgia (E.T. Wherry s.n. PH). Various sources 
have attributed South Carolina to the range of G. brevifolia (NatureServe Ex- 
plorer 2005; South Carolina Plant Atlas 2005; USDA 2005; Weakley 2005), based 
upon specimens from Clarendon (J.F Townsend 384 CLEMS), Lee (C.A. Aulbach- 
Smith 4070 and 4077 with S. Hutto USCH), and York counties {f.B. Nelson 821] 
with L. Lundquist USCH), South Carolina, but these specimens, however, have 
been annotated by either Deborah Lewis or the second author as G. ramosa 
(Clarendon County) or G. viscidula (Lee and York counties). 

The nativity of discoveries, such as this, are highly debatable, and entire 
papers have been devoted to the topic (Lamont & Young 2005). We consider G. 
brevifolia a native component of the flora of Delaware, the Delmarva, and the 
Mid-Atlantic region for several reasons. First, though the habitat supporting 
the species in Delaware is highly degraded, G. brevifolia is found in similar habi- 
tats in other portions of its range. Second, two other species of Gratiola, G. 
ramosa and G. viscidula, once reached their northern range limits on the 
Delmarva Peninsula. The historic Maryland population of G. ramosa was also 
significantly disjunct, ca. 500 km (300 mi), from its closest known occurrence 
in Hoke Co., North Carolina (Radford et al. 1968). Third, the genus Gratiold is 
known for having strange and highly disjunct distributional patterns. For in- 
stance, De Lange (1997) reported G. pedunculata R.Br. as a new addition to the 


828 BRIT.ORG/SIDA 22(1) 


flora of New Zealand and considered the species to be a recent natural migrant 
likely distributed by vagrant birds from mainland Australia, a disjunction of 
ca. 2150 km (1332 mi). Furthermore, G. virginiana, a species primarily distrib- 
uted in the southeastern United States, is disjunct to the states of Veracruz and 
Puebla, Mexico from central Texas, a distance of more than 900 km (525 mi) 
where it occurs in association with other species characteristic of the temper- 
ate southeastern United States (Miranda & Sharp 1950). Even G. brevifolia has 
a fragmented range characterized by wide disjunctions. For example, on the 
Gulf Coastal Plain, the easternmost population of G. brevifolia west of the Mis- 
sissippi River in Rapides Parish, Louisiana is separated by ca. 623 km (387 mi) 
from the westernmost population east of the Mississippi River in Pike Co., Ala- 
bama. Lastly, G. brevifolia is not known to be cultivated or grown for horticul- 
tural purposes; therefore, it is highly unlikely t 
introduced from nearby cultivated plants. 


a 


rat this species would have been 


ACKNOWLEDGMENTS 

Appreciation is extended to several people who contributed information or com- 
ments to this project: Curtis Hansen, Auburn University; Frank Hirst; James 
Macklin, Philadelphia Academy of Natural History; Bill McAvoy, Delaware 
Natural Heritage Program; Lucile McCook, University of Mississippi; Robert 
Naczi, Delaware State University; John Nelson, University of South Carolina; 
Heather Sullivan, Mississippi Department of Wildlife, Fisheries and Parks; Chris 
Frye of the Maryland Department of Natural Resources, Wildlife and Heritage 
Service; and Ron Wilson. Additional thanks are given to the following herbaria 
where specimens were examined: AUA, BRIT, CLEMS, DOV, LSU, MISS, NCU, 
NO, PH, TENN, USCH. 


REFERENCES 
Brown, M.L.and R.G. Brown. 1984. Herbaceous plants of Maryland. University of Maryland, 
College Park. 
CHesrer, E.W., B.E. Worrorp, and R. Krat. 1997. Atlas of Tennessee vascular plants. Volume 2. 
Angiosperms: dicots. Misc. Publ. No. 13. The Center for Field Biology, Austin Peay State 
University, Clarksville, TN. 


De Lance, PJ. 1997. Gratiola pedunculata (Scrophulariaceae): a new addition to the New 

Zealand flora. New Zealand J. Bot. 35:317-322. 

FerNatb, M.L. 1950. Gray’s manual of botany (ed. 8). Dioscorides Press, Portland, OR. 

GteAson, H.A. and A. Cronouist. 1991. Manual of vascular plants of northeastern United 
States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx. 

Goprrey R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United 
States: Dichotyledons. Univ. of Georgia Press, Athens. 

Lamont, E.E.and S.M.Youns. 2005. Juncus diffusissi an addition to the flora of New York, 
with notes on its recent spread in the United States. J. Torrey Bot. Soc. 132:635-643. 


KNAPP AND ESTES 829 


McAvoy, W.A. and K.A. Bennett. 2001. The flora of Delaware: an annotated checklist. Dela- 
ware Natural Heritage Program, Delaware Division of Fish and Wildlife (Document No. 
40-05/01/01), Smyrna, DE. 

McAvoy, W.A. 2003. Rare vascular plants of Delaware. Delaware Natural Heritage Program, 
Delaware Div. of Fish and Wildlife, Smyrna, DE, unpublished report. 

Miranpa, F.and A.J. SHare. 1950. Characteristics of the vegetation in certain temperate re- 
gions of eastern Mexico. Ecology 31:313-333. 

NatureServe Explorer: An online encyclopedia of life [web application]. 2005. Version3.1. 
NatureServe Arlington, Virginia. Available .org/explorer.(Accessed: 17 
Dec 2005). 

Pennett, EW. 1935.The Scrophulariaceae of Eastern Temperate North America.Wickersham 
Printing Company. Lancaster, PA. Pp. 78-80 

Plank, M.O.and WS. ScHENck. 1998, Delaware Piedmont Geology. Delaware Geological Sur- 
vey, University of Delaware, Newark, Delaware, Special Publication No.20, 69 pages. 

Raproro, A.E., H.E. AHLEs, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. 
University of North Carolina Press, Chapel Hill, NC. 

Sorrie, B.A.and A.S.Weakiey. 2001. Coastal plain vascular plant endemics: Phytogeographic 
patterns. Castanea 66:50-82. 

SouTH CAROLINA PLANT ATLAS. 2005. University of South Carolina. Available http:// 
cricket.biol.sc.edu/herb/. 

Tatnatt, R.R. 1946, Flora of Delaware and the Eastern Shore. Soc. Nat. Hist. of Delaware. 
Lancaster, PA. 

Weaktey, A.S. 2005. Flora of the Carolinas, Virginia, and Georgia, Working Draft: June 10, 
2005. Available: http:// herbarium.unc.edu 

USDA (United States Department of Agriculture). 2005. Natural Resources Conservation 
Service. The PLANTS database (plants.usda.gov/plants/). National Plant Data Center, 
Baton Rouge Louisiana (Accessed: 17 Dec 2005). 


830 BRIT.ORG/SIDA 22(1) 


Book Notices 
University of California 


MELINDA A. Zeper, DANIEL G. BRADLEY, Eve EmsHwitter, and Bruce D. Smitu. 2006. 
Documenting Domestication: New Genetic and Archaeological Paradigms. 
(ISBN 978-0-520-24638-6, hbk.). University of California Press, California/ 
Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, 
US.A.(Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $70.00, 
361 pp., b/w photos, illustrations, 8 1/2" x 11" 

The book is divided into four sections: 1) Archaeological Documentation of Plant Domestication. 2) 

Genetic Documentation of Plant Domestication. 3) Archaeological Documentation of Animal Do- 

mestication. 4) Genetic Documentation of Animal Domestication. 

From the Publisher—Agriculture is the lever with which humans transformed the earth over 
the 


the last 10,000 years and created new lorms of plant and animal species that have forever alterec 
face of the planet. In the last decade, significant technological and methodological advances in both 


molecular biology and archaeology have revolutionized the study of plant and animal domestica- 
tion and are reshaping our understanding of the transition from foraging to farming, one of the ma- 
a turning g points in human history. This groundbreaking volume for the first time brings together 


eading veologists and biologists working on the domestication of both plants ve ani uc to 
pee a wide variety of archaeological and genetic approaches to tracing the 
of domesticates. It provides a comprehensive overview of the state of the art in this ene cals 
recent findings on specific crop and livestock species in the Americas, 


field as well as reviews of 1 
Africa. Offering a unique global perspective, it explores common challenges and poten- 


Eurasia, anc 
tial avenues for future progress in documenting domestication. 


M. Nevin Smit. 2006. Native Treasures: Gardening with the Plants of Califor- 
nia. ISBN 0-520-24425-7, pbk.). University of California Press, California/ 
Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, 
US.A.(Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $24.95, 
278 pp., color photos, 7" x 10" 

From the Publisher—The author explains how California's diverse terrain, climate, and geology sup- 

port a wealth of plant species—more than 6000—and offers suggestions for designing with most ol 


cultivation, as well as with some more obscure but garden-worthy groups. With 
sity celebrates 


= 


the major né 
an engaging narrative and a wealth of illustrations, this ode to beauty and dive 
-alifornia’s rich store of native plants and encourages readers to visit them in rn native haunts 


and invite them into their gardens. 


SIDA 22(1): 830. 2006 


HEDYOTIS AUSTRALIS (RUBIACEAE) NEW TO MISSOURI 
AND: FLORIDA AND RELATED SPECIES IN THE 
SOUTEL-CENTRAL UNITED STATES 


Walter H. Lewis 


Department of Biology 
Washington University 
St. Louts, Missouri 63130, U.S.A. 


ABSTRACT 


The distribution of white-flowered Hedyotis australis is extended to Missouri and Florida as well as 
within Alabama, Arkansas, Mississippi, and Tennessee. This distribution is compared to allied 
homostylous species H. crassifolia and H. rosea which also flower as winter annuals in the south- 
central United States. Hedyotis crassifolia is a diploid species having a flowering peak earlier than 
the tetraploid H. australis. An example of aaa out- eee within H. crassifolia via bee pollina- 


=a) 


tion is discussed. Hedyotis rosea is now ¢ ‘tirpated in Missouri 

RESUMEN 
Hedyoti tralisde fl bl distribuida por Missouri y Florida asi como también 
en Alabama, Arkansas, Mississippi, y Tennessee. Su distribucion se copa la de las otras especies 
homostilas H. oS ia y H. rosea la region centro- 
sur de Estados Unido ide Hedyoti ifolia ti le f] 


5 
— 


a especie ene H. aera Se Sale un emul de posible entrecruzamiento dente de 
H.c ae via polinizacion por abejas. H ra extinguida en Missouri. 


INTRODUCTION 


Three winter annual and homostylous bluets (Hedyotis or Houstonia) flower 
and fruit in the south-central U.S. largely between January and April. The most 
common and widely distributed of these is the purple- or violet-colored Hedyotis 
crassifolia Raf. (Houstonia pusilla Schépf). Less well-known is a smaller an- 
nual with tiny white corollas which is easily overlooked even when flowering 
in the same habitat and at the same time as H. crassifolia. Consequently, this 
southern white-flowered Hedyotis australis WH. Lewis & D.M. Moore 
[Houstonia micrantha (Shinners) Terrell] is infrequently collected even when 
common. A third very small bluet with large pink- or rose-colored corollas, 
Hedyotis rosea Raf.{Houstonia rosea (Raf.) Terrell], is more narrowly distributed 
and occurs in eastern Texas, Louisiana, central and southern Arkansas, west- 
ern Mississippi, and southeastern Oklahoma, with outlying populations in Ala- 
bama (Tuscaloosa Co.) and in 1931 southeastern Missouri (Dunklin Co.). 
During March 2005, collections were made of these species concentrating 
on areas where H. australis was unreported but could be expected to occur. Find- 


SIDA 22(1): 831 — 836. 2006 


832 BRIT.ORG/SIDA 22(1) 


ing these small bluets by their distinct flower colors of purple-violet, white, 
and pink-rose is one of the best ways to identify them. In addition, corolla tubes 
of H. australis are short with calyx lobes as long as or longer than the tubes 
(Fig. 1), while lobes of H. crassifolia are much shorter than its longer tubes. How- 
ever, without these floral characters their discovery is greatly limited even when 
growing side by side as they often do, and particularly since anthesis is con- 
fined toa few weeks at most. All collections were made by Walter H. Lewis and 
Memory Elvin-Lewis; their numbers and herbaria of deposit are provided for 
each collection cited. Many additional ones are at MO. 

Cytological differences also exist between these species: H. crassifolia is 
diploid, 2n = 2x = 16, x = 8, H. australis is tetraploid, 2n = 4x = 32, x = 8, and H. 
rosea is diploid, 2n = 2x = 14, x =7,a reduced base number from the others. That 
eos a uctions, and to some extent polyploidy, have played impor- 
tant roles in the evolution of these and other North American species of Hedyotis 
(Lewis 1965) is supported by recent molecular phylogenetic analyses by Church 
(2003) who ei found that “Houstonia is not distinct from either North 
American Hedyotis or Stenaria.” Additional research is needed lor North Ameri- 
can (including Mexican) taxa and others worldwide before generic- and tribal- 
level complexes of both the Hedyotideae and Spermacoceae are satisfactorily 
resolved. Clearly, this paper does not provide additional data regarding classili- 
cation and phylogeny, for it is intended only to extend certain species distribu- 
tions and comment briefly on specific ecologic and reproductive observations. 

To aid the reader in identifying the three species, as well as two forms of H. 
crassifolia, f. crassifolia and [. albiflora (Standley) W.H. Lewis, characters were 
obtained from Lewis (1970), Smith (988), Terrell (1996), and herbarium collec- 
tions at BRIT and MO: 


— 


1. Corollas white, lobes and tube each 1.5-2.5 mm long; calyx lobes as long as or 


longer than corolla tube (Fig. 1) australis 
1. Corollas purple-violet or pink-rose, rarely white, tube exceeding length of calyx lobes. 
2. Stems 3 eles cm long; corollas purple-violet, occasionally pale purple to white, 
lobes 2.5-3.5 mm long, tube 3.5-4.5 mm long; calyx lobes about half as long as 
corolla eee nmon ____ crassifolia 
3. Corollas purple-violet throughout f. crassifolia 
3. Corolla lobes white to pale purple-violet, tube purple-violet to reddish 
f. albiflora 
2. Stems 1-3 cm long, often clumped; corollas pink-rose fading to oe yes tol 
nearly white lobes 2.5-4.5 mm long, tube 5—7 mm long; calyx lobes 1.2-2.5 mm 


long; occasional rosea 


Hedyotis australis 

Prior to 2005 the known continuous distribution of H. australis was limited to 
eastern Texas, Louisiana, Arkansas (excluding the northeast), southeastern 
Oklahoma (only McCurtain Co.), western and central Mississippi, and south- 
western Tennessee (only Shelby Co.), in addition to outlying populations in cen 


LEWIS AND FLORIDA 833 


Fic. 1. Hedvoti tralis W.H. Lewis & D.M.M inl f t y, Natchitoches, N hitoches Parish, Loui- 
siana, WH, Lewis 15867 (MOQ).A h g calyx lot long Ila tube ( imes longer). Bar equivalent to 
about 1 cm 


tral Alabama (Tuscaloosa Co.), and central and eastern Georgia (Bibb and Co- 
lumbia Co.’s) (Chambers 1965, Lewis 1968, Terrell 1996). Its current distribu- 
tion is now extended to include: 

(1) southeastern Missouri.—northern Dunklin Co., N of Campbell, 15968 F 
MO, UMO, US; northern Dunklin Co., NW of Campbell, 15969 MO; Butler Co., 
Poplar Bluff, 15970 MO, US. These are the first reports for Missouri. The Poplar 
Bluff collection at 36'76'N is the most northerly record of the species, although 
this distribution is similar to the new record at Paris, Tennessee (36'17’N). 
Hedyotis australis was found in Missouri’s Bootheel region and adjacent Butler 
County in low foothills at the southeastern edge of Ozark Plateau where a 
three collections grew in grassy cemeteries and where herbicides were not used. 
None were found in nearby pastures or meadows or in lower-lying areas where 
cotton is now grown in large farms and where herbicides are widely used. The 
species was also absent from flood-plains and other riparian habitats to the east 
along the Mississippi River valley. 

(2) northeastern Arkansas.—Greene Co., Paragould, 15921 MO, 15964 BRIT, 


834 BRIT.ORG/SIDA 22(1) 


MO. As in Missouri H. australis was not found in the Mississippi River valley, 
but only along a low ridge just west of the Missouri Bootheel among grass in a 
hilly cemetery. This is its most northeasterly location in Arkansas which par- 
allels its northwestern distribution (Smith 1988). 

(3) western panhandle of Florida.—Escambia Co., Hwy 4 at Sandy Hollow 
Rd, 15936 FSU, MO, USE This new outlying record was found in a hilly region 
of extreme northwestern Florida growing with H. crassifolia in grass by a road- 
side corner. Both species were in flower 22 March, although flowering had just 
begun for H. australis (no mature capsules) compared with numerous flowers 
and maturing capsules of H. crassifolia. Tetraploids like H. australis often have 
peak flowering later than their diploid relatives (as H. crassifolia) and this gen- 
eralization applies here (Lewis 1980). Hedyotis australis was not seen elsewhere 
in Escambia County, adjacent Santa Rosa County, or nearby Mississippi and 
Alabama, although H. crassifolia was found with regularity. 

(4) eastern Mississippi.-Hinds Co., Hwy 27 at Bear Creek Rd, 15930 BRIT, 
MO; Smith Co., Taylorsville, 15932 MISS, MO, US; Itawamba Co., Fulton, 15950 
DAO, MO; Lee Co., Tupilo, 15951 MISS, MO; Union Co., New Albany, 15953 MO, 
US; Marshall Co., Holly Springs, 15954 MISS, MO. In Mississippi's southeastern 
area H. australis was only occasionally found along grassy roadside verges, but 
where found it was common while H. crassifolia was less so (Hinds, Smith Co.’s); 
in the northeast it was widespread in the more frequent upland areas (Itawamba 
Co., the most easterly Mississippi collection, Lee, Union, Marshall Co.’s) where 
H. crassifolia was also less common. Observation of fewer flowering plants sug- 
gests that {lowering of H. crassifolia had peaked some time earlier rather than 
limited frequencies of plants. 

(5) western Alabama.—Marion Co, Hamilton, 15949 BRIT, MO, US. Plants 
of this second collection known for Alabama were growing in abundance among 
other invaders in lawns. This new extension eastward in Alabama from those 
found in nearby northeastern Mississippi is about 80 miles northwest of the 
only other known Alabama collection at Tuscaloosa in Tuscaloosa Co. (Terrell 
1996) where H. australis was re-collected (15945 BRIT, MO, US) growing with H. 
crassifolia 15946 BRIT, MO) and nearby (ca. 20 m) H. rosea 15947 BRIT, MO, 
US), the most easterly record for this species. 

(6) western Lennessee.—Tipton Co., Covington, 15926 MO; Fayette Co., Hwy 
57 at Hwy 18, 15956 MO, US; Hardeman Co., near Bolivar City, 15957 MO, TENN; 
Gibson Co., Humboldt 15960 MO, TENN, Bradford 15961 MO, TENN; Henry Co., 
Paris, 15962 MO, TENN. Tothe one collection of H. australis previously reported 
in the extreme southwestern part of the state at Memphis (Shelby Co.) (Terrell 
1996) are added numerous collections throughout western Tennessee as far east 
as 88° 15°W in Fayette, Gibson, Hardeman, Henry, and Tipton Counties found in 
lawns, hillsides, ruderal areas, and cemeteries. As in Missouri, the species was 
not found in the Mississippi River valley, but did occur with some frequency on 


LEWIS. 835 


the eastern ridge and uplands eastward and as far north as near the Kentucky 
state line to Mississippi in the south. Although the species was not found in 
western Kentucky, it might occur there. 


Hedyotis crassifolia 

Many collections were made of H. crassifolia whose range, including H. minima 
(Beck) Torrey & Gray, extends from Maryland to Georgia west to Wisconsin 
and Iowa, and south from Kansas to central Texas along the Gulf Coast states to 
the panhandle of Florida (Terrell 1996). Often it occurs with H. australis and 
less commonly also with H. rosea, particularly in eastern Texas and Louisiana 
where all three species can be found in flower together. All are homostylous 
with different chromosome numbers; recently Church (2003) found H. 
crassifolia with cleistogamous flowers among greenhouse-grown plants. In- 
breeding is clearly predominant and no hybrids between the three species have 
been found. Thus, I was surprised to observe foraging in a population of H. 
crassifolia a bee, Agapostemon virescens (Halictidae), extending its proboscis 
into the corolla tube while wings beat and rapidly flying from one flower to 
another after a few seconds stay. It visited about 30 flowers (occasionally re- 
turning to the same one) of 12 or so plants during 3-4 minutes and then flew 
off. This occurred near the Appalachian ridge in Alabama, Clay Co., at the Bap- 
tist Church cemetery adjacent Highway 49, 23 March. No other species of 
Hedyotis was found in the vicinity; in fact few species were in flower. How com- 
mon such visits might be in the early spring (few pollinators seen) is unknown, 
but that it was observed even once suggests a means of at least limited out- 
crossing for this homostylous species. One could venture that its distributional 
success covering a wide range of habitats compared to other related winter an- 
nuals might be due to the success of even secondary out-crossing in combina- 
tion with in-breeding via homostyly and possibly cleistogamy. 


Hedyotis rosea 

In representative specimens for H. rosea Terrell (1996) includes a collection from 
Missouri in Dunklin Co., near Campbell, 5 April 1931, collected by A.L. Grant 
s.n.(MO). Recently George Yatskievych located a second sheet collected by Julian 
Steyermark 406 (MO) on the same day, both undoubtedly made during the same 
collecting trip. Steyermark’s label locality is more precise at “ca. 5 miles north- 
west of Campbell” in open flat ground surrounded by Quercus phellos woods. 
This is the same area of northern Dunklin Co. where new records of H. austra- 
lis were found only in “protected” (from herbicides) cemeteries on 28 March 
2005. Searches around Steyermark’s locality, in much of the northern part of 
the county, and in neighboring counties proved fruitless. I suggest that this 
single outlying population collected in 1931 is extinct and that H. rosea is now 
extirpated from the Missouri flora. No collection has been found in neighbor- 


836 BRIT.ORG/SIDA 22(1) 


ing Arkansas or Tennessee and the nearest H. rosea in central Arkansas (Terrell 
1996) is about 175 miles southwest of the original Missouri find. Indeed the loss 
of habitat and heavy use of herbicides for many years in the Bootheel counties 
may have contributed to the loss of H. rosea and perhaps other species whose 
ranges once extended to this region. 


ACKNOWLEDGMENTS 
| thank Memory Elvin-Lewis, Washington University, who assisted me in the 
field during both 2005 collecting trips, Richard Clinebell, entomologist with 
the Missouri Botanical Garden, who determined the bee discussed under 
Hedyotis crassifolia, and N. Rogerio Castro for translating the Abstract into 
Spanish. lalso acknowledge the use of important collections of Hedyotis at BRIT 
and MO. 


REFERENCES 


CHameers, K.L. 1963. Hedyotis australis in Georgia. Rhodora 65:271-273. 

CHurcH, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneup- 
loidy and breeding system evolution in the radiation of the lineage across North 
America. Molec. Phylogenetic Evol. 27:223-238. 

Lewis, W.H. 1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia 
(Rubiaceae). Amer. J. Bot. 52:257-264. 

Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 
50188) 

Lewis, W.H. 1970. Hedyotis L.In:D.S. Correll and M.C. Johnston. Manual of the vascular plants 
of Texas, Texas Research Foundation, Renner. Pp. 1487-1490. 

Lewis, W.H. 1980. Polyploidy in species populations. In W.H. Lewis, ed. Polyploidy: Biological 
relevance. Plenum Press, New York. Pp. 103-147. 

SuitH, E.B. 1988.An atlas and annotated list of the vascular plants of Arkansas, ed. 2. Pub- 
lished by the author, Fayettesville, AR. 

TerreLt, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48. 
The American Society of Plant Taxonomists. 


Book REVIEWS 


TosHio YosHipa. 2005. Himalayan Plants Illustrated Hi hokubutsu daizukan 
(ISBN 4-635-58031-8, pbk.). YAMA-KEI Publishers, Co., Ltd., 1-9-13 Akasaka, 
Minato-ku, Tokyo, JAPAN. Price: ¥13,000 (ca. $115), 800 pp., (773 pp.in color), 
18.5 x 26 cm, in Japanese. 


When I returned from field work in Asia in September of 2005, a book of beautiful photographs of 
sree wildflowers was among the items in Ge mail. The book, Himalayan Plants Illustrated, is 


by Toshio Yoshida, one of the most talented, knowledgeable and intrepid photographers of alpine 
Bee in eastern ee ae Yoshida has traveled ne ly to capture images of the characteristic aes 
of the Greater Himalayan region. Having also visited some of the areas where he has worked, I wa 
amazed to see that he | g p eray of the species in one of the world ee 
areas of plant pages 

The photog bool he f nine auEnOrs = ia of field studies in the Hima- 
] di China and from leadi The photos are richly 
su upplemen tedb iled f h dy of herbarium specimens and fror 


the literature. The introductory material inc ludes a fresh discussion of floristic zones and their de- 
limitation in the Greater Himalaya, southern Xizang (Tibet) and the Hengduan Mountain region of 
China, based on the author’s observations and interpretation. 

The nt contains 2,739 color photos covering 1,771 plant taxa, of which 1,586 are at species 


rank, 40 are subspecies, 61 are varieties, 2 are forms, and the remaining 82 are unidentified. The pho- 
tos were taken in all parts of the Himalayan range; northern Pakistan, Kashmir, northwestern India, 
— Sikkim, Bhutan and Tibet, as discussed in more detail in the front pages of the book. The book 


compared with Flowers of the Himalayas by Oleg Polunin and Adam Stainton, published in 
1964 by Oxford University Press. Polunin one Stainton’ s book covers 1495 plant taxa, with 694 color 
1 in the last 128 pages, and 3 gs. Because of the politics of those days, the 


field sites covered by Polunin and Stainton are in about half the Himalayan range, excluding Bhutan, 
Tibet and most parts of Pakistan and Sikkim 
n Mr. Yoshida’s book the photos and descriptions/discussions are on the same page, crossed 
Gus by number. Plants exhibiting a wide range of variation, a likely phenomenon in the 
malaya, are represented by two or more photos to give a sense of eens variation. Below 
a photo is the plant name in Japanese, which is often the Japanese transliteration of the scientific 
me. Also preva if bce is ou icient space, are details of the photograph, Hues date, place, 
i pie and i the eine The letters A-Z3 preceding the place 
name below each photograph ee to the 34 maps on the front pages. A brief guide to the 
ee is on the inside flap of the sae ine and can be read without turning pages 
ach photograph is keyed to : nding text entry. The text entry gives the Japanese 
name, the scientific name and synonyms, if anys fe Sliawed ee flowering period, altitudi- 
nal distribution and details of the habitat and life form. The morphological description in the text 
matches the plants in the corresponding photo(s), since it is basically derived from author's field 
notes at the time the photo was taken and from dried specimens of the photographed a? sif they 
were obtained. The oa are further enriched with information from the literature. 
The 82 Dice meine’ cen which may represent new species or varieties, also text with 
data about th ts. They are provided with the name of the species that the author 


} bs ] 


result Del C1 

Plant morphology, especially of species dwelling in the alpine zone, is strongly affected by the 
environment. With that thought i in mind, the author has tried to include photos that show not only 
the plant, but its habit and its place within the habitat. 


SIDA 22(1): 837. 2006 


838 BRIT.ORG/SIDA 22(1) 


Che introductory material includes articles on the history of research on Himalayan plants 
written by Prof. Hideaki Ohba on pages 4-7, geobotany of the Himalaya on pages 8-1], and a map of 
the whole Himalaya. On pages 12-25 are 34 topographical maps. Botanical guides to the areas of the 
aya are on pages 26-32. An analysis of the horizontal and alecidinal: distribution patterns of 


—_ 


Hima 
the plants covered in the book is on pages 33-39 and adaptive strategies of alpine plants in the 
aya are covered on oo 40-43. Himalayan plants in the oe mountains of southwest 


—_— 


Hima 
China are discussed on pages 
though the book is ae the high quality and detail in the photos make the diversity 
of the flora and vegetation in this unique and highly diverse area easy to comprehend. The number 
of individual taxa included for each genus provides a good sense of the range . shel within 
large genera. Some examples: Arisaema, 17; Astragalus, 24; Gentiana, 33; Impatiens, 22; Leontopodium, 
10; Meconopsis, 21; Pedicularis, 66; Rhododendron, 58; Saxifraga, 06; Sdussurea, 40. 
book is beautifully bound on high quality paper and one of the nice features is t 
pages lie flat when the book is opened at any page. Rumor has it that an English language edition is 
being considered.—David E. Boufford, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, 
MA 02138-2020, U.S.A, david_ boufford@harvard.edu. 


rat the 


Q. 


Two British Orchid Books 


ANNE and Simon Harrap. 2005. Orchids of Britain and Ireland: A field and site 
guide. (ISBN 0-7136-6956-X, pbk., durable field guide binding). A & C Black, 
38 Soho Square, London, W1D 3HB, U.K. (Orders: +44 (0)20 7758 0200 
customerservices@acblack.com; http://www.acblack.com/search.asp) 
£29.99 ($51.95 US), 480 pp, 360 color plates, numerous watercolors, line art, 
49 maps, glossary, bibliography, index, 21.6 x 13.8.cm. 

MicHaet Fotey and Sipney Clarke. 2005. Orchids of the British Isles. SBN 0-9541916- 
1-7, hbk.). Griffin Press Publishing Limited, 35 Wessex Way, Cox Green, 
Maidenhead, SLO 3BP, UK (Orders: http://www.griffinpress.co.uk/ 
orchids.htm). £45 ($78 US), 390 pp., extensively illustrated with color 
photographs; 50 maps, anaes bibliography, index, 27.6 x 20.5 cm. 


It is not often a reviewer has t] ity to review two eae that cover the same subject 
and are pubis! hedat the same time. eThisis the situation with both books on the orchids of the British 
Isles. Each | from the British Isles, Both books are packed with full- 
color photographs, maps, eeebnical details, and similar information. Yet they are as different from 
her as apples and oranges. While Foley and Clarke’s large format book (published in associa- 

tion the Royal Botanic Garden, Edinburgh), contains the more technical information, the Harraps’ 
work is designed as a field guide—and that it very much is. Asa field guide the binding is a durable 

Flexibind type that is sown and open tat at any given pom Is eal) ita Pabst binding i is an injus- 


dare \ ry detail. A 


i= 
a 


tice. The photographs in Orchids of Britain 
though both books use the maps oe Ne Atlas of the British and Irish Flora ioe: reproduced in 
the Harraps’ book are much easier to read owing to the fact that the distributional dots representing 
county records for specific time segments are in red, black, and green as opposed to those in Foley 
and Clark that are in graduated shades from black to grey to red to pale pink. The wealth of informa- 
tion that appears in both books is very complete and covers every possible aspect of the orchids in- 

cluding their on, W ae extensive notes on more than 0 supspccics, varieties, and forms; dis- 
tributional i and status. $ although ina slightly different 
format in each book. In Orchids ide and Ireland the Harraps go even further and detail th 


SIDA 22(1): 838. 2006 


BOOK REVIEWS 839 


history of the known populations and their current status. Both books treat the genera Dactylorhiza 
and Epipactis in great detail and make every effort to sort out the species, subspecies, and varieties— 
no simple task! It is not surprising that the bibliography and glossary are similar in each work. 

Given that both books cover much of the same information what does one book have that the 
other does not? 


Foley and Clarke @ £45 Harrap @ £29.99 
covers 55 ee covers 56 species 
historical information extensive specific information for seeking the 
superior chapter on orchid taxonomy orchids (44 page site guide to 330+ places to 
checklist of species see orchids 
authors names with the species easier to read distribution ma 
photographic hints chart for each species ae eine over the 


keys to the genera and species of selected genera years 
chapters by guest authors on cultivation and — convenient field guide format 
conservation growth pattern gr ae of several genera 
comparative plate of the bee orchids, Ophrys, of 
Europe 


Both books use the term variety is the sense of forma—color and growth forms and both are cio of 
the misuse of the term albino. A true albino plant = no oe other than white-stems, leave 
and flowers. What is usually meant is white-fl fte hthe name albiflora. aioe 
it is the only publication of the two with keys for identification, be and Clarke is by no means a 
field guide and the size limits its use as such. Having a copy of both books would be ideal. Use the 
Harraps in the field and then when returning home consult gens and cl arke for confirming or addi- 
ae information. The photographs in both books at, but in Foley and Clarke many of 

‘ull-page images have blurred. I feel certain the er ee were sharp but they were en 

ee just a ou beyond a limits. The me) momceavie enor found in HOY and Clans The first, 
under Spi simply piranthes 
{x}steigeri being a hybrid ee i 5 romanzoffiana and S. cernua. It is not been viewed as such for 

over 25 years. Spiranthes {x}steigeri is a synonym for S. ochroleuca. Information under Pseudorchis 
albida stating that it occurs in North America from the east to Alaska is incorrect. The species, as P. 
straminea, is restricted to northwestern Newfoundland and a single site in western Quebec on the 
shore of Hudson's Bay. Both facts could have been easily found in Flora of North America. 

ographical information is given for both Foley and Clarke and their guest authors but none 

for the ieee It would have been nice to know a bit acon: them and their other work. All four 
ae authors should be congratulated on their worl contribution 
to the 21" Century's knowledge of wild orchid in the British Isles and are heartily recommend d fo 
both i: novice and Byorsionala alike. If youare visiting the British Isles for an orchid hunting expe- 
dition the An Simon Harraps'’ field guide will be especially useful —Paul Martin Brown, au- 
thor, Wild meee ‘of Florida, The Wild Orchids of North America, Wild Orchids of the Southeastern 
United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. naorchid@aol.com. 


5; 


rt 
— 


Orchids of Mexico 
E. HAcsater, M.A. Soto ARENAS, G.A. SALAZAR CHAVEZ, R. JIMENEZ MACHORRO, M.A. 
Lopez Rosas. and RL. Dresser. 2005. Orchids of Mexico. (ISBN 9-6878-8908- 
X, hbk.). Published by Instituto Chinoin, A.C. Mexico City. Spanish and English 
versions available. (Orders: redactamex@yahoo.com.mx; Redacta, S.A. de 


SIDA 22(1): 839. 2006 


840 BRIT.ORG/SIDA 22(1 


CV, Avenida 10 de Mayo # 249, Col. San Pedro de los Pinos, C.P.03800 Mexico, 
D.F MEXICO). $100.00, plus shipping, 304 pp., 650 color photographs, 13.25" 
mR IO(3I 9 R25 5c) 


Initial disappointment may often give way to surprised delight. Such is the case of Orchids of Mexico. 


Given the publication and research history of the estimable authors | had expected a detail taxo- 
nomic oe the orchids of Mexico. Imagine my surprise when a large format-33.5 x 25.5cm- 


oO 


‘coffee table’ book arrived! This sumptuously illustrated volume is much more orchids ‘in Mexico’ 
than ‘of Mexico’ and wastes no time in luring the reader to that country. | soon discovered that the 
| a searchable CD that will cover 


the 164 genera and 1200+ species with over 1500 color photogr aphe a checklist, and cross-referenced 


taxonomic treatment | was so looking forward to is fort 


synonymy. The current publication mentions 162 of those genera and 450 of the species. 
But back to the volume at hand; sitting before me is one of the lushest books on orchids | have 
ever seen. Apart from the text, which we will get to later, it is lavishly illustrated with the highest 
quality piptogs ae and es ae oduction I have seen in many years and the printing on heavy 


| 
=p 


stock only y of the book. The authors have provided us with not just an overview of 


the oi tha nat grow w eh Mexico, but with details of the several climatic and geological regions 


of the country shown at first in a two-page layout of a satellite image of Mexico. After relating the 
history of orchids, and orchid research in Mexico, with many reproductions of plates from well- 
known historic volumes, it concludes with the ethobotanical history featuring Vanilla planifolia. A 
brief chapter on orchid biology is fae illustrated with photographs of plants from flower to fruit 
to flower with the many stages in between. 

ow to our adventure searching ne wilds of Mexico. As we visit the eight various habitats the 


narrative highlights the many species of orchids that are present and and numerous vies of the habi- 
tats and illustrates them with color nes Sai At this a few synonyms are given and occa- 


sionally the reader may be temporarily confused with some of the newer g c concepts for famil- 


iar species such as Prosthechea citrina for Cililes: citrinad and Tamayorkis for a few species 


traditionally known as Malaxis. Many species found in Mexico are also found in the bordering re- 
gions of the southwestern United States, Texas, and Florida. It is always welcome to see photographs 


of these more southerly sire that may be very rare in the United States. Because of the arrange- 


ment of chapters by habitat it req sulting the index to find information on specific species. 


But that is not the apparent aim of this book. It is to immerse the reader in the diversity of the orchids 
of Mexico and the verdant and prolific variec 


habitats. A final chapter on Conservation efforts and 


successes is followed by detailed photo creits and an extensive bibliography. 


es 


nks to the team of authors and photographers the text is scientifically accurate and this 
reviewer did not note any mipurectly ae deled or positioned graphics (a feat not many books of this 


scope can boast!). There few i with a missing letter Le, seudobulb for pseudobulb, 
but nothing that could | essibly distract? from the intense text. Many photographers were credited in 


the book and the meticulous editing of these photographs has provided the finished book with a 
continuity not often seen in similar publications. 

Is it expensive?—yes, at $100—but worth every penny of the price. Keep in mind that two books 
were really written; one in Spanish and then one in English. Were Orchids of Mexico not sponsored 
by Chinoin lam sure it would have to sell for much more or not even have ever gone into production. 
If you have any interest in North American ore ae Finca rom the southern regions of that 

rl} t . 1 


s volume. And remember, that the interac- 
tive CD will soon be available to satisfy those a us sas thrive on taxonomic treatments!.—Paul Mar- 
tin Brown, author, Wild Orchids of Florida, The Wild Orchids of North America, Wild Orchid sof the 
Southeastern United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. naorchid@aol.com. 


O 
‘e) 
— 
~ 
=> 
= 
=) 
io) 
+) 
la 
} 
S 
1) 
= 
a | 
s 
y 
gy 
i= 
C 
= 
7 
~_ 
iS 
i 
a 
£ 
= 
7 


SIDA22( (1): 840. 2006 


BOOK REVIEWS 841 


Orchids of Cuba 


Juan LLAMACHO (text) and JuLio A. LARRAMENDI (photographs). 2005. The Orchids 

Cuba—Las Orquideas de Cuba. (Cloth. ISBN 84-933615-2-6, hbk.). Greta 

see Plaza Mayor, 24 25340 VERDU, SPAIN. (Orders: +34 690 733 O11, 

http://www.gretaeditores.com inlo@gretaeditores.com). +9.00 Euros [$58.19 

2/13/06], 287 pp., Spanish-English in parallel columns, color photographs, 
interpretive graphics, table of species, and glossary, 9.6" x 7" 


Following a recent trend for ae orchid books to be both scientifically accurate and written 
such a manner that they are popular with the non-academic enthusiast, The Orchids of Cuba fulfills 
a geographic niche ee has ie been void. This work of two enthusiastic Cubans has produced not 
only the first full-color book on the orchids of the island archipelago but the first text in English 
(Spanish/English in parallel columns) that provides an alluring enticement to the island nation. 
Following an imprimatur by Carlyle A. Luer, M.D. and foreword by James Ackerman, Ph.D. the books 
starts with the basic biology of orchids and biogeography of Cuba and proceeds to merge the two, 
describing and illustrating the several regions ol the r nation. These ieee and the orchids found 
within them are then treated in detail. Of the 305 sr ted from Cuba at the writing of 
the book, more than 150 are treated in detail and, for the most spate have excellent color photographs. 
Both the author's and photographer's love of the pleurothallids (Lepanthes and related genera) is 
me in the spectacular close-up photographs of many of these tiny species. Following the species 
treatments is a section on undescribed (those areas that have not yet been identified and/or de- 


cribed to science) species, with the same high-quality photographs. The book cana: witha table 
of all 305 ly found in Cuba, their flowering times an g n,a 
brief biniosoeny: and an ‘excellent glossary. What is unfortunately ae isan ee Because of 
the arrangement of species it makes it difficult to find s ickly. A listing of syn- 
onyms for the species treated in full Id also have been nihelpfa ul. No oe to ee are given 
as this book is, ey just the prelude to more publications covering all of the orchids of Cuba. 
F 


e reviewer's standpoint The Orchids of Cuba has the same proble her works 


that treat orchids from the Caribbean and Central America/Mexico. W ee Ee are stated the 
occurrence a the Spaces in the United States, usually Florida, is omitted. In part, tk € 
in several works. Rather than Ph: on pation 


fr rom a quarter century or more ago, simply consulting Flora of North America or Wild Orchids of 
Florida would have solved this problem. Those species included in es or and also documented 
lata, Bulbophyllum 


Florida include lonopsis utriculariodes, Prosthechea boothiana,S 
Pe ee Epidendrum nocturnum, Liparis nervosa, Maxillaria crassifolia, Brassia caudata, 
Eulophia elata, Malaxis pane neal distans, Phaius tank eae avert! amo ee 
drum radicans, Prescottia opligantha, and Pelexia adnata. Govenia trib- 
uted to Florida (Govenia floridana is - correct eae From a taxonomic standpoint ane issue 
can be taken other than to note that the treatment and photographs of Habenaria quinqueseta are 
oe H. ues yn. H. quinqueseta var. macroceratits) ane the curious treatment of 
Sa 


inceolata and Stenor hy nchos squamulosum. The a tw | ld be treated as Sacoila 
ee - 


| I lif { | Sacoila and Steno 


or absence of a mentum (in the latter). 

The English translation is excellent and very readable and the only error noted would be the 
incomplete name, or omission of a period after the abbreviation, in one of the authors’ names: Gal for 
Galeotti. The book is pamiede on very cies Blew heavy paper and durably bound. It is to be highly 
recommended toall i f the Caribbean and south Florida as well.—Paul Mar- 


SIDA 22(1): 841. 2006 


BRIT.ORG/SIDA 22(1) 


tin Brown, author, Wild Orchids of Florida, The Wild Orchids of North America, i d oe rids of the 
Southeastern United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. na 


Splendor in the Hill Country Grass 

BRIAN and Suir_ey Lorin. 2006. Grasses of the Texas Hill Country. 2006. (ISBN 
1-58544-467-7, flexbound with flaps). Texas A&M University Press, 4354 
TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 979-458-3982, 979- 
847-8752 fax; http://www.tamuedu/upress/). $23.00, 208 pp., 170 color 
photos, 7 lin art, 2 maps, black and white figures, glossary, bibliography, 
index, 53/4" x 81/2". 

Grasses of the Texas Hill Country is a pictorial guide to grasses of the Hill Country region of Texas. 

Designed for the layperson, this book is full of crisp color photographs on a black background that 

clearly show the inflorescences (seed heads) of grasses common in the Hill Country as well as many 

other areas of Texas. Authors Brian and Shirley Loflin have created a guide that is easy to use with 

grasses grouped by pan in inflorescence structure. The front and back book flaps serve as 

ler types and the icons used throughout the text. 


handy 
The ope ag chapter of the book introduces readers to the Hill Country region and other veg- 
etation regions of Texas. The introduction discusses the basic structures 5 of grass and includes color 
d t e different inflores- 


images. Within the “how to use this book” section is a detail 
cence structure types used to organize the grasses in the book. This ene section includes 

other items of interest such as a discussion on how . are ey scientific names, the meaning of 
those names, and a list of both the genus and commo es of grasses included in this book. 

here are seventy-seven grasses included in the or Each grass has an associated profile that 


eo 


includes features of the species, plant uses, growing season information, and habitat information in- 
cluding soil, location and distribution. The features section of each profile includes important infor- 

mation such as typical height, description of grass inflorescence, some vegetation characteristics, 
whether the grass is cool or warm season, native or introduced and whether the grass is annual or 
perennial. Much of the feature information is included in a bar under the common name of the grass 

ina rather intuitive icon form. The economic usage sections describe a species’ use as a forage grass, 
pace grass, aneseane grass, or value tow beanies ce grass profile has at least one large image of 
the int photographs of growth habit in nature. 


There are a few items that would have made this book more useful to both beginners and more 
experienced grass speenva tnete are ® fantastic pee ar as of the inflorescences included with 
each grass, but unfortur ately pho tographs of the vegetative structur ies (ligule 
and auricle) needed to nee identify grasses when the inflorescence is not available. However, the au- 
nin the “features” of the plant descrip- 


an 


thors have included some vegetative structure information wit 
tion. It would have also been beneficial to have the growth habit photographs th lat accompany many of 


the plant descriptions for every grass included in the text. There were a couple of typos/mix-ups no- 
ticed; an example is the aioe is p hoa in Figure 5 where the auricle was incorrectly labeled 
as the ligule. Some other small t roticed. Despite these few items, there is no doubt that this is 


a very useable book for persons with an interest in learning grasses of the Hill Country area 
If you are a beginner to grass identification or are looking for a good color photograph guide to 
grasses tor the Hill Country (and surrounding area) of Texas, Grasses of the Texas Hill Country by 
Brian and Shirley Loflin would be a terrific book to start with. The glossary, writing style and icon 
quick guide add to make te ee ial eos an easy to reference and use. This book ise sa) recom- 
of: 


mended for use b planti cation enthusiasts.—Lee Lucke ydoo, 


*y 
Herbarium, Botanical Research Thsti tute afie xas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(1): 842. 2006 


BOOK REVIEWS 


FREDERICK CAMPBELL and RICHARD Dube. 1997. Landscaping Makes Cents. (Paper- 
back Edition). (ISBN 0-88266-948-6.). Storey Publishing, 210 MASS MoCA 
Way, North Adams, MA, 01247. U.S.A. (Orders: # 66948, 1-800-441-5700, 
wwwstorey.com). $16.95, 170 pp., color and b/w figures, appendices, glos- 
sary, further reading guide, index, 81/2" x 11" 


If you want to improve the look of your homes’ landscape and increase its’ resale value at the same 
time, pe sure to pick ap a copy of i Landscaping Makes Cents yy ceapod and Dube. The book is 
| who is either Ul landscape by themselves or with 

the help of an outside contractor. The book is div ided ne three main sections, the first of which isa 
discussion of how landscaping can add or detract value from a home. The second section is how to 
plan for hardscape elements and plants as well as on thoughts on HOuRUIBUnE a — The final 
section contains all the info you need to know when hiring an 
The first section is subdivided into four chapters that can help readers recognize which ele- 

ments of ier a add phe) most value to EheDE property; these ee include several tip boxes 
pecial | tics. Some of the more specific topics include how to 


deteumine the values of trees, pl “int igen removal, adaition, environmental value, historic value 
and aesthetic value. The final chapter of the first section s] addresses landscape consider- 
ations when buying, puree and sell ing a home. 


ine second section of tl 
hi I hei dit land- 


id e anninga pices: anda landscape 


scaping considerations for ine with various periods of time until oe ele and finally 
the idea of financing help. Readers will need to investigate into budgets for their own situations, as 
the inlomtation included i in cl hapter five on budgeting is ie generalized. Chapter six of this sec- 
as well as multiple assessment tools to help readers think through and de- 
sign their ee For eck one of the first tools er in this chapter is “What do you 
want in your landscape?” with seventy-plus | uae epien: you ee want, including patios, stor- 
hie ies S. ane authors 


and an 


age sheds, small fruit trees, hot tubs, benches, gaze 
melee a short discussion of aesthetic landscape He p ve lities for 
outline for plotting the actual design. Chapter seven discusses investing in cee elements with 
inclusion of wall types fences patios, we alkways arbors drip edges ee and others. Once again, 
the authors include a number of helpful tip and eonsideradon boxes. Chapter eight provides the reader 
ideas for planting in their landscape. There are discussions of different plant types, growth forms and 
some helpful hints and te for planting plants. One would need to do further research into the 
m well in their ea Hula areas (the Anno’ nae provided a Usbe2 zone al 

_Th fj lch rot 


plants that would ae 
diness map inan 
scape. Readers are provided with many aoe and ideas for planting a Suse able Saas mee 
an excellent assessment tool for performing a self-environmental audit of ones’ own rty. 

The final section of the book contains three chapters, which cover all aspects of hiring a pro- 
ee ce. contractor. For those readers interested in hiring such a person, these chapters 
will be especially helpful for you. This section's first chapter discusses how to hire a contractor be- 
ginning with the titles, definition and services offered by various landscape professionals; it includes 
things to ask about prior to hiring the person. The second chapter of this section focuses on negotiat- 
chapter is a very in-depth chapter on what questions to ask of contractor 


ing contracts. The final 
during the planning sae and how to Rian specific concerns. There is also an included appendix 
on how to evaluate the aesthetic value of ones’ landscape. 
ee ng Ma i Cents by Frederick Campbell and Richard Dube is easy to read and includes 
helpful tip eeu aes cies uae topics for nae in planning and designing a home land- 
Lands ng as a fantastic resource for readers who have an in- 


SIDA 22(1): 843. 2006 


844 BRIT.ORG/SIDA 22(1) 


Es 


terest in, either personally or through a landscape contractor, designing their home's landscape to 
improve both its financial and aesthetic value.—Lee Lucke ate - rbarium, Botanical Research In- 
stitute of lexas, 509 Pecan Street, Fort Worth, TX 76102-4060, U. 


Ropert H. MOHLENBROCK. 2006. Aquatic and Standing Water Plants of the Cen- 
tral Midwest: Filicineae, Gymnospermae and Other Monocots, Excluding 
Cyperaceae. (ISBN 0-8093-2670-1, hbk.). Southern Hlinois University Press, 
Carbondale, IL, U.S.A. (Orders: 618-453-6633, http://wwwsiuedu/‘siupress). 
$65.00, +00 pages, b&w line drawings, glossary and indexes to the com- 
mon names and genus and species, 6" X 9", 


Aquatic and Standing Water Plants of the Central Midwest: Filicineae, Gymnospermae and other 
monocots, excluding Cyperaceae by Mohlenbrock is a flora, which consists of the ferns, conifers, and 
other monocots, excluding the sedges portion of the larger Aquatic and Standing Water Plants of the 
Central Midwest series. The Cyperaceae family is included in another book by the same author within 
the same series title. The book’s Central Midwest range includes: Ohio, Indiana, Kentucky, Illinois, 


lowa, Missouri, Nebraska, and Kansas 

The flora includes information on 221 species, including members of the genera Azolla, 
Wood wardid, Equisetum, lsoetes, Lycopodiella, Osmunda, Acorus, Sagittaria, Elodea, Iris, Juncus, Lemna, 
Najas, Platanthera, Spiranthes, Calamagrostis, Echinochloa, Glyceria, Leersia, Zizania, Heteranthera 
Potamogeton, Sparganium, Lypha, Xyris, and others. There is no overall key to ie mine family, but 


= 


the individual families have keys to determine genus, as well as within genus keys to determine spe- 


cies. Each species is presented in the book by black and white drawings, which provides images in- 
cluding: plant habit, leaves, inflorescence, achene, fruit, and/or sheath/ligules a many grasses, as 


las an accompanying detailed description. 
he description for each species contains a great deal of information, including the current 


we 


accepted specific epithet, synonyms, as well as the authority and references for publications on each 


specific epithet. Each species’ description includes plant habit information, descriptions and mea- 
surements of leaves and flowers (when applicable), root types, inflorescence, notable colors and, where 
ap] plicable, details on parts unique to the family or genus, such as root collar information in Poaceae. 


The description eeecases information, witha listing of which states the species have wetland 
designations (ie., FACW) within the U.S. Fish and Wildlife Wetlands Inventory. The author has also 


included handy oe field traits within the plant description to hel p determine that species from 


similar looking others. 

Aquatic and Standing Water Plantsof the Central Midwest: Filicineae, Gymnospermae and other 
monocots, excluding Cyperaceae by Mohlenbrock would be a nice addition to the library of any per- 
ee Mohlenbrock’s book on Cyperaceae within the 


son who works with aquatic plants, right a 
same series. Although this series focuses on the Midwest, a number of the species included in this 
flora are found in various regions of North America. The black and white drawings are crisp, easy to 
read images that show major Saran traits for the species. Some familiarity with family traits 


will be necessary to know which family key to use to identify a plant. The keys are straightforward, 
but the flora does contain a glossary for any unfamiliar terms, and is user friendly. The description 
for each species is thorough, which can help you conclude that you have determined the correct spe- 


] ] 
cies or not; and includes very helpful “in the field” identification hints. If you need to identify mono- 


cots and ferns in aquatic regions, be sure to pick up this helpful book. —Lee Luckeydoo, Herbarium, 
Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(1): 844. 2006 


BOOK REVIEWS 845 


French/English Review 


Dov FE Sax, JOHN J. StAcHOWwicz, and STEVEN D. Gaines (eds.). 2005. Les invasions 
despéces: Considérées dans le cadre de l’écologie, de Pévolution et de la 
Biogéographie. Sinauer Associates, 23 Plumtree Rd, Sunderland »MA oe 
0407, US.A.(Orders: 413-549-1118 fax; ord om; Ww 
$51.95, 495 pp., b&w figures and tables, 7"x 91/4". 


ee —Ce sate constitue une vue a ensemble de Hompueuet> pECHEICH ayant été menées sur les 


éc ologique | 1 l’évolution 
a ws x a o >) a L | ee 


hinl 


De sept chapitres ont leurs peopes illustrés grace a des exemples tirés de publications 
i bien le milieu aquatique, aérien que 


E 

eae 
es grands noms de la recherche se sont rassemblés pour mettre leur savoir en commun. La 
contribution ee ae Soles a peums de prendre en considoauog un large éventail de eal 
6 AUX iaires, en passant par le role des 


maladies infectieuses sur les communautés ace on exemple. 
rés les auteurs, ce livre peut-étre lu dans sa globalité afin de connaitre les tenants et 
aboutissants de nombreuses études antérieures, mais peut également étre abordé par chapitre 
same selon les préoccupations de chacun. 
e lecteur peut Come Nae e comment Petude des espéces invasives est devenue une discipline 


res dé s. Par exehipis Petude des BSpeces | invasives permet 


sient phare depuis | 


‘appréhender la facon dont : ver les pro- 


cessus en temps réel, plutét que de déduire les processus ayant eu lieu dans | e passé seulement grace 
aux modéles quils ont pu engendré 

Ce livre est le dernier ouvrage sorti et constitue le premier de référence pour toute personne 
voulant orienter ses recherches vers les théories les plus récentes et vers les méthodologies dediées a 
létude des invasions d’espéces. 


English.—This book is an overview of previous research as well as current investigations about spe- 
cies invasions. The study of non-native mere their invasions can provide insights into ecology, 
evolution, and biogeography. 

ere are seventeen chapters that cover everything from — to aerial and terrestria 


inva- 


sions; each chapter ends with its own set of supporting references. Leading researchers were brought 
together to review the field of invasion Pioloay. an contri bution of 45 scientists has provided the 
editors with a diverse array of i communities, avian populations 
on islands, to the role of the infectious diseases on natural communities. The editors note, “Individu- 
ally, we believe that each of these chapters has something significant to offer. Collectively, we ho 
that this book has much to offer to both invasion biology and to our fundamental understanding _ 


ecology, evolution, and biogeography.” 
reader wil 


— 


quickly gain an understanding of why the study of invasive species has be- 
come an important scientific discipline in the last few decades. For instance, the study of invasive 


ae helps to better understand the function of an ecosystem. “Looking closely at invasions allow 
s to observe processes in real time, rather than having to infer the operation of processes that oc- 

see in vihe past solely from the aus they eave 
s the most recent book on species invasions and is recommend it to anyone inter isis in 


the ee of invasive species . isa source o oD to 
odologies and will t king in this discipli Virginie 


SIDA 22(1): 845. 2006 


846 BRIT.ORG/SIDA 22(1) 


H. Raquet, Graduate Student, Botany Curatorial Assistant, ELL. d Herbarium Department of Bio- 
logical Sciences Texas Tech University Flint & Main ae nae , TX 79410-3131, U.S.A 


vireinie raquet@ttuedu 
Oo tT 


ANNA Pavorb. 2005. The Naming of Names. (ISBN 1-59691-071-2; hbk.). Bloomsbury 
Publishing, 175 Fifth Ave., New York, NY 10010, U.S.A. (Orders: http:// 
www.bloomsbur y.com/). $45.00, 384 pp., illustrated, 8" x 9 5/8" 


T 
animals, germs, stars, storms, rocks, and other huge kingdoms have eventually ee broken down 
into types and grouped so that we could begin to understand them. The Naming of Names traces t 


— 


ae natural world presents innumerable objects which humans have needed t i | 


he 
search for order in the natural world. Such a process has taken many centuries, and we have gotten 


better at it with a scientific understanding of the world, but the impulse has been there for as long as 
we have been thinking about the things around us. Pavord, author of the The Tulip (2001), Flower 
Power: the Meaning of Flowers in Art (2003), and an expert gardener, details the history of plant tax- 
onomy from the ancient Greeks to 17th-century British botanist John Ray. She reveals the history of 
plant classification and shows how the process was affected by intellectual, political and cultural 
cues The journey, traced here in detail for the first time, involves the culture of Islam, the first 


the Indies and the first eons in the New World. 
In Athens, Aristotle's pupil Theophrastus was the first man ever to write a book about plants. How 
can we name, sort, and order them? He asked. 7 The lebat s still, two thousand years later. The 


Naming of Names gives a compelling nsigne intoa wen in oF intr igus and patensely sonrpeHtye egos. 
She has gone back to the ancient Greeks 


sance tried to get a grasp on the disorderly a kingdom, with eventual success even bel fore the taxo- 
nomic standards see down by Linnaeus which we still follow. It is the pre-Li in efforts that Pavord 


has chr icled. 1 reisalsoa histo OTY O of plant illustration W ithin the se pages. The eventual woodcuts did 


not have to Be = with many meprodiices Here S107 ae swirling masses of plants or delicate leaves in 


fine detail. 1 t books could show enough useful detail to 
be excellent field g ike dithaush for centuries authors neliea on previous works of folklore. 

Along with Theophrastus, Pavord’s highest praise goes to Englishman John Ray, who in 1696 coined 
the term “botany.” He provided six rules by which to categorize plants, not only the ones familiar to 
him in England, but the spectacular finds being brought from distant lands. Others had previously 
insisted on classifying plants by use, which was entirely artificial, or more helpfully by leaf or seed 
form, but it was Ray who ue pokes on its ou irst real nec oi nouns the distinction of seeds that 

sprout with one leaf ort ). He 


kne W he Was pal 18 ol 
an ongoing process, predicting hae future botanists would look back and “our proudest discoveries 
will seem slight, obvious, almost worthless.” He might have been right, but seen as a tribute to their 
efforts, Fhe Naming of Names shows how fe discoveries, achieved over eas nine ao curious, ia 
voted, and fallible plantsmen, have brought us to our current und 


ane ith Ray, barely mentioning the recent advances that have been ee with DNA testing; euch 
ests have confirmed much of what was eventually realized as o speauae es tree, but have upset 
ac Pere as well. It nas peeH a a long botanical trip 


5», and P. ‘ ps shi 1 inclusion of 


rmous fun. The book eee illustrated, with a third of 
the | pages being taken up with ileaniees (most in color) nicely keyed to the text 
Int 


he best sections, she slows down to draw detailed portraits of researchers and describe how 
each contributed to the ay evolv ing sand until Me ate 1000s, Enmenee) science of botany. The 
] 


story makes 


] { 1] | } 
Lue i HIAaAGE LIC ee 


Ol 


comprehensible. It would be an eelledt choice for Sublic libraries w ‘ith al ipi sted in 


the history of botany.—Gary Jennings, Botanical Research Institute of Texas, 509 ee Street, Fort 
Worth, TX 76102-4068, U.S.A. 


SIDA 22(1): 846. 2006 


an - 1d i in si A ais - 7 


; 


JUL TANA | BAM ae 1 ( yay * el i, rey Pub A ANS fA! PUORIDA TOVVA ee 


gS A 7. VIRGINIA OREGON ‘NE -BRASK eS A NEW Me NICO. C 
J PUPP? ORTH CAROLINA SOUTH CAROLINA MONTAR 


Vv olf the Hlustrated Flora of East Texas is the first fully 
illustrated flora for East Texas, a species-rich area that 
contains roughly two-thirds of all the plant species known for 
lexas. The book covers all the native and naturalized ferns and 
similar plants, gymnosperms, and monocotyledons (1,060 
species) known to occur in East Texas 

Published by Botanical Research Institute of Texas 

Pars NG Texas Floras Pro ojec 


£7, 


proj Texas (B 
and the 

Austin College Center for Environmental Studies. 
Pr f| 


od 


x10 //" 
4 pages 
1; ee bw sia 5 
] ,0004 
Sida, oe 426 Illustrated Flora 
es of East Texas 
isbn 1-889878-12-X, X, hbk. 
George Diggs, Barney Lipscomb, 
May 2006 Monique Reed, Robert J. O’Kennon 
$89.95 Texas residents add 8.25% tax ($7.42) From Big “D" 


' ' of beau prone iDisecken we groves of timber, 
please inquire) ( 


rchids nay ads, azaleas, and Texas Pride! 


Available from Botanical Research Institute of Texas Press 
50 


9 Pecan Street 
Fort Worth, TX 76102-4060, U.S.A ‘I 


E-mail: sida@brit.org 


Fax: 1-8 2-4112 Fost: AO 


www.brit. Lae a/sbm/ 


BRIT 


his photographic guide to the Wild Flowers of 
; - ; WILD FLOWERS OF MOMBACHO 
Mombacho is color-coded and arranged by Family NICARAGUA 


within the color sections. It covers flowering plants found Sie alee ee MOM 


on the whole of the Volcano; which includes both tropical 
cloud forest and dry, deciduous forest, at lower altitudes. 
The book covers 200 plants, each of which has two or 
three photographs and a short description in both English 
and Spanish giving common name(s); a short description 
of the plant; height; flowering period; habitat and any 
known use of the plant. The vast majority of plants are 


likely to be found in similar habitats throughout tropical HELEN PICKERING 


America and nearly one third are pan tropical, making this 


book of interest to a wide audience throughout the tropics. WILD FLOWERS OF MOMBACHO 
Sida, Bot. Misc. No 28, 2006 pisiieck 
issn 0823-1475 FLORES SILVESTRES DEL MOMBACHO 
isbn 1-889878-14-6 BY HELEN PICKERING 
6.5” x 8.5", x + 217pp. 
over 500 photographs @ 
$15 + p&p" 
‘International: $7 BI 


211 
Texas residents add $1.24 sales tax PRESS 


: RRANSAS FLORIDA IOWA MiISSISS 


Barney L. Lipscoms, EDIToR 
Botanical Research Institute of Texas 
509 Pecan Street 


Fort Worth, Texas 76102-4060, USA 
2-444] / 817 332-4112 FAX 


Electronic mail: sida@brit.org 


Home page at the URL: http://www.brit.org/sida/ 
S1DA CONTRIBUTIONS TO 


BOTANY WAS FOUNDED BY 7 JOHN W. THIERET, ASSOCIATE EDITOR 


Lovo H. SHINNERS (LEFT) Dept. of Biological Sciences 
Northern Kentucky Universit 
IN 1962. INHERITED BY Perel epee ea Ta 

Highland Heights, KY 41099 USA 
WILLIAM F. MAHLER 


(RIGHT), DiRecTOR EMERITUS FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR 


oF BRIT IN 1971, AND Dpto. de Botanica, Facultad de Biologia 


l 


niversidad de Leon 

SINCE 1993, IT HAS BEEN 
E-2471 Leon, SPAIN 

PUBLISHED BY Brit PRESS. . os 
The views expressed in this journal do not necessarily 


ate reflect those of the editors or of the Botanica 
Research Institute of Texas 
Guidelines for contributors are available upon request 
and on our SIDA home page as well as the last pages 
of each volume. 


Subscriptions for year 2006: 
$39. Individual 

$75.USA Institutions 

$85. Outside USA 


numbers issued twice a year 


OSIDA, CONTRIBUTIONS TO BOTANY 


VOLUME 22, NUMBER 2, PAGES 847-1274, 


COPYRIGHT 2006 


BOTANICAL RESEARCH INSTITUTE OF TEXAS 


PRINTED IN THE UNITED STATES OF AMERICA 


SSN 0036-1488 


TABLE OF CONTENTS 


A new name for Sida, Contributions to Botany (1962-2006) 

BARNEY LipscomB 

SYSTEMATICS 

A new species of witch-hazel (Hamamelis: Hamamelidaceae) apparently 
endemic to southern Mississippi 

S.W. LEONARD 

Eriogonum callistum (Polygonaceae), a new species from the Tehachapi 
Mountains of California 

JAMES L. REVEAL 

Six new combinations in Dodecatheon (Primulaceae) 

JAMES L. REVEAL 

Ericameria ubsp phila (Asteraceae), a new rabbitbrush from 
the San Luis Valley of Colorado 

LorAN C. ANDERSON 


A new species of Boltonia (Asteraceae) from the Ridge and Valley physi- 
ographic province, U.S.A. 

JOHN F TOWNSEND AND VESNA KARAMAN-CASTRO 

A new species of waterlily (Nymphaea minuta: Nymphaeaceae) from 
Madagascar 

KENNETH LANDON, RICHARD A. EDWARDS, AND P. IVAN NOZAIC 


Two new combinations in Peyritschia (Poaceae: Pooideae: Aveninae) 
Victor L. FINOT, PAUL M. PETERSON, AND FERNANDO O. ZULOAGA 


Poa matri-occidentalis (Poaceae: Pooideae: Poeae: Poinae), a new species from 
Mexico 

PAuL M. PETERSON, ROBERT J. SORENG, AND YOLANDA HERRERA ARRIETA 

Bromus ayacuchensis (Poaceae: Pooideae: Bromeae), a new species from Peru, 
witha key to Bromus in Peru 

JEFFERY M. SAARELA, PAUL M. PETERSON, AND NANCY F. REFULIO-RODRIGUEZ 
Psidium cauliflorum (Myrtaceae), a new species from Bahia, Brazil 

Lesiie R. LANDRUM AND MARCOS SOBRAL 

New species of Myrtaceae from Ecuador and Peru 

Bruce K. Hoist AND MariA LUCIA KAWASAKI 

Zeuxine pantlingii, sp. nov. (Orchidaceae), a new species from India 

AVISHEK BHATTACHARJEE AND HJ. CHOWDHERY 

Four novelties and a lectotypification in Matelea (Apocynaceae: 
Asclepiadoideae) from Hispaniola 

ALEXANDER KRINGS 

New combinations in the genus Cymopterus (Apiaceae) of the southwestern 
United States 

RONALD L. HARTMAN 


873 


Six new species of Sphagnum (Bryophyta: Sphagnaceae) from North America 
RICHARD E. ANDRUS 


Review of Crataegus series Pulcherrimae (Rosaceae) 

J.B. Puipes, RJ. OTKENNON, AND K.A. Dvorsky 

Crataegus series Bracteatae and Triflorae (Rosaceae) 

J.B. Pipes, R. LANCE, AND K.A. Dvorsky 

Morphometric analysis of an Amelanchier (Rosaceae: Maloideae) complex 
on the Delmarva Peninsula (Delaware, Maryland and eastern Virginia) re- 
solves the taxonomic identities of Amelanchier obovalis and A. canadensis 
CHRISTOPHER T. FRYE 


Carex reznicekii, a new widespread species of Carex section Acrocystis 
(Cyperaceae) from eastern North America 

Davip A. WERIER 

A new species of Eugenia (Myrtaceae) from the Monteverde region, Costa 
Rica 

FRreD R. BARRIE 

Taxonomic review of Symphyotrichum patens (Asteraceae: Astereae) 

Guy L. Nesom 

Drepanostachyum falcatum var. sengteeanum identity and origins (Poaceae: 
Bambusoideae 

C.M.A. STAPLETON 

Rejection of lectotypification of Aster amellus (Asteraceae: Ast ) and se- 
lection of a new lectotype 

JOHN C. SEMPLE 


ANATOMY AND MORPHOLOGY 

Trichome morphology in selected Mexican red oak species (Quercus section 
Lobatae) 

M. Lucia VAZQUEZ 

BOTANICAL HISTORY 

Thomas Walter Typification Project, I. Observations on the John Fraser folio 
DANIEL B. WarD 

FLORISTICS, ECOLOGY, AND CONSERVATION 

Evidence for hybridization between two sympatric violet species, Viola 
grahamii and V. hookeriana (Violaceae), in central Mexico 

AuREA C. CorTES-PALOMEC AND Harvey E. BALLARD, JR. 

Herbivory of feral goats on Espiritu Santo Island, Gulf of California, Mexico 
José Luis LFON-pDE LA LUZ AND REYMUNDO DOMINGUEZ-CADENA 


A demographic and ecological analysis of Geocarpon minimum 
(Caryophyllaceae): a federally threatened species in southwest Missouri 
TIM SMITH AND JOSEPH S. ELy 


1009 


1027 


1049 


1071 


1075 


1081 


1087 


1091 


1111 


1119 


1135 


1145 


A quantitative study of styles and achenes of terminal and basal flowers of 
Schoenoplectus hallii (Cyperaceae), a rare plant species of transient wetland 
habitats 

MaRrIAN SMITH, SARA AMMANN, NANCY PARKER, AND PAIGE METTLER-CHERRY 


Annotated checklist of the vascular flora of the Lance Rosier Unit, of the Big 
Thicket National Preserve, Hardin County, Texas 

Larry E. BROWN, BARBARA R. MACRopserts, MICHAEL H. MACROBERTS, 

PauL A. HARCOMBE, WARREN W. PRuEss, I. SANDRA ELSIK, AND 

STANLEY D. JONES 


An updated, annotated vascular flora of Caddo Parish, Louisiana, with notes 
on regional phytogeography and ecology 

BARBARA R. MACROBERTS AND MICHAEL H. MACROBERTS 

Carex stipata (Cyperaceae), Chamaesyce geyeri (Euphorbiaceae), Eurytaenia 
texana (Apiaceae), Pediomelum esculentum (Fabaceae), and Talinum 
calycinum (Portulacaceae) deleted from the Louisiana flora 

BARBARA R. MACROoBERTS AND MICHAEL H. MACROBERTS 


New vascular plant records for New Mexico 
RONALD L. HARTMAN, BRIAN REIF, B.E. NELSON, AND BRIAN JACOBS 


We . (Cc 1 7 
f£ 


Galium tricornutum (Rubiaceae) and P iaceae) 
new to Oklahoma 


Amy K. BUTHOD AND BRucE W. HOAGLAND 


Capraria mexicana (Scrophulariaceae) in Cameron County, Texas: rediscov- 
ered in the United States 
ALFRED RICHARDSON AND KEN KING 


Range expansion of Hexalectris grandiflora (Orchidaceae) in Texas 
MARGARET BROWN-MARSDEN AND ANNE B. COLLINS 

Erodium malacoides (Geraniaceae), new to Texas 

Davip E. LEMKE AND JED L. APLACA 

New distribution records for Eupatorieae (Asteraceae) in the United States 
Guy L. Nesom AND BILLIE L. TURNER 


1159 


1175 


1191 


1221 


1225 


1235 


1237 


1239 


1245 


1249 


Book reviews and notices 862, 866, 894, 904, 930, 954, 958, 1008, 1026, 1074, 1090, 1134, 


1144, 1158, 1174, 1190, 1220, 1224, 1234, 1254 
Announcements 847, 1236 


Sida, Contributions to Botany update—1257 
Reviewers for Volume 22, 2006-1258 
Index to volume 22, 2006 
Titles of Articles With Authors—1260 
Authors—1264 
Botanical Names and Subject—1266 
New Names and New Combinations—1272 


Index to new names and new combinations in Sida 22(2), 2006 


Boltonia montana J.F Townsend & V. Karaman-Castro, sp. nov—874 

Bromus ayacuchensis Saarela & P.M. Peterson, sp. nov—919 

Calyptranthes manuensis B. Holst @ M.L. Kawasaki, sp. nov—931 

Carex reznicekii Werier, sp. nov—1052 

Cymopterus breviradiatus (WL. Theob. & CC. Tseng) R.L. Hartm., comb. & stat. nov— 
955 


Cymopterus davidsonii (J.M. Coult. & Rose) R.L. Hartm., comb. nov—956 
Cymopterus glomeratus var. fendleri (A. Gray) R.L. Hartm., comb. & stat. nov—956 
Cymopterus sessiliflorus (WL. Theob. & CC. Tseng) R.L. Hartm., comb. nov—956 
Dodecatheon clevelandii var. gracile (Greene) Reveal, comb. nov—863 
Dodecatheon clevelandii var. insulare (HJ. Thomps.) Reveal, comb. nov—863 
Dodecatheon clevelandii var. patulum (Kuntze) Reveal, comb. nov—863 
Dodecatheon pulchellum var. macrocarpum (A. Gray) Reveal, comb. nov—863 
Dodecatheon pulchellum var. shoshonense (A. Nelson) Reveal, comb. nov—864 
Dodecatheon utahense (N.H. Holmgren) Reveal, stat. nov—864 
Ericameria nauseosa subsp. ammophila L.C. Anderson, subsp. nov—868 
Eriogonum callistum Reveal, sp. nov—857 
Eriogonum Sect. Lanocephala Reveal, sect. nov—857 
Eugenia haberi Barrie, sp. nov—1071 
Eugenia yasuniana B. Holst @ M.L. Kawasaki, sp. nov—934 
Hamamelis ovalis S.W. Leonard, sp. nov—850 
Matelea corynephora Krings, sp. nov—942 
Matelea phainops Krings, sp. nov—948 
Matelea rhynchocephala Krings, sp. nov—949 
Matelea torulosa Krings, sp. nov—951 
Nymphaea minuta K. Landon, R.A. Edwards & PI. Nozaic, sp. nov—887 
Peyritschia howellii (Hitchc.) Finot & PM. Peterson, comb. nov—897 
Peyritschia pinetorum (Swallen) Finot & P.M. Peterson, comb. nov—899 
Poa matri- occidentalis P.M. Peter: son & pOune sp. nov.—906 

oa matri lentalis sub I i g & PM. Peterson, subsp. nov—911 
Psidium cauliflorum Landrum & Sobral, sp. nov—927 
Sphagnum beothuk Andrus, sp. nov—966 
Sphagnum bergianum Andrus, sp. nov—964 
Sphagnum kenaiense Andrus, sp. nov—961 
Sphagnum mcqueenii Andrus, sp. nov—959 
Sphagnum sitchense Andrus, sp. nov.— 969 
Sphagnum talbotianum Andrus, sp. nov—970 
Zeuxine pantlingii Av. Bhattacharjee @ HJ. Chowdhery, sp. nov—935 


A NEW NAME FOR SIDA, CONTRIBUTIONS TO BOTANY 
(1962-2006) 
Barney Lipscomb 
Botanical Research Institute of Texas 
Fort Worth, Texas 76102-4060, U.S.A. 
barney@brit.org 


Beginning in 2007, the journal Sida, Contributions to Botany will no longer be published 
by that name but instead will be called the Journal of Botanical Research Institute of Texas 
(J. Bot. Res. Inst. Texas). The new journal will have a new look but generally the same format, 
botanical focus, and editorial direction will remain. 


Reason for chang 

The leadership of. the Botanical Research Institute of Texas has decided the name “Sida” 
is offensive and has too much of a negative connotation because of its similarity to the 
word meaning AIDS in Spanish, French, Italian, and Portuguese. Perhaps the name change 
is a good thing but lam personally saddened to see Shinners’ original and long-standing 
name displaced. 


Brief History of Sida 

1962—Lloyd H. Shinners, a member of the Southern Methodist University faculty anda 
prolific researcher and writer who wanted to edit his work and the work of others, pub- 
lished the first issues of Sida. He named the journal for a genus of yellow-flowered plants 
of the mallow family, distributed throughout the world and especially common in Texas. 
Shinners continued as editor and publisher until his death in 1971. 

1971—Wnm. FE. Mahler, professor of botany at SMU, became editor and publisher. 
1977—Barney Lipscomb became assistant editor to Wm. F Mahler. 

1982—Barney Lipscomb became editor, Wm. F Mahler remained publisher. 
1993—With Barney Lipscomb as editor, Sida became a publication of the BRIT Press. 
2002—BRIT celebrated the 40th anniversary of Sida, Contributions to Botany. 
2006—BRIT Press publishes last issue of Sida, Contributions to Botany. 


Thank you 

To all authors, we thank you for choosing Sida, Contributions to Botany. A big thank you to 

the hundreds and hundreds of individuals who generously supported Sida through their 

time and expertise in reviewing more than 1000 manuscripts published from 1962- ZOO 
Thanks to all authors, reviewers, subscribers (individuals, institutions, 

and readers for your continued interest and support. It is our plan to continue bringing 

you the best sources of current research in classical and modern systematics through the 

Journal of Botanical Research Institute of Texas with your continued support. I personally 

wish you the best in 2007 and beyond. Your comments are welcome. 


SIDA 22(2): 847 — 848. 2006 


848 


Publication dates of individual issues of Sida, Contributions to Botany. 


BRIT.ORG/SIDA 22(2) 


Volume Issue Pagination Publication Date Volume Issue Pagination Publication Date 
22 2 847-1274 22 Nov 2006 10 4 269-341 27 Nov 1984 
oy) 1-846 11 Aug 2006 10 3 195-267 8 Jun 1984 
21 4 1955-2511 21 Dec 2005 10 2 95-193 28 Nov 1983 
21 3 1249-1954 5 Aug 2005 19 1-93-28 Apr 1983 
21 2 511-1248 28 Dec 2004 9 4 269-381 7 Dec 1982 
21 1 1-150 20 Aug 2004 9 3 195-268 10 Jun 1982 
20 4 1343-1836 22 Dec 2003 2 2 “194 24 Nov 1981 
20 3 877-1341 29 Jul 2003 = : 1-97-29 Apr 1981 
20 2 461-876 23 Nov 2002 8 4 317-372 17 Dec 1980 
20 1 1-460 19 Jul 2002 8 3 218-315 11 Jun 1980 
19 4 767-1233 28 Dec 2001 8 2 123-216 30 Nov 1979 
19 3 445-766 23 Aug 2001 : ep eee Tee 
19 D 235-444 20 Dec 2000 7 4 309-397. 13 Dec 1978 
19 1 1-234 23 Sep 2000 7 3 227-308 30 Jun 1978 
18 4 941-1316 23 Dec 1999 d 2 94-226 9 Dec 1977 
18 : 631-940 20 Sep 1999 if 1-93 27 May 1977 
18 2 377-630 28 Dec 1998 6 4 243-331 29. Sep 1976 
18 1-376 15 Oct 1998 6 3 123-242 14 May 1976 
17 4 645-873 31 Dec 1997 6 2 63-122 11 Nev 1375 
17 3 533-643 12 Sep 1997 S ee “eaaee 
17 2 305-532 30 Dec 1996 5 5 301-361 28 Feb 1975 
17 1-303 30 Sep 1996 5 4 191-300 16 Sep 1974 
16 4 613-809 29 Dec 1995 5 3 137-190 21 Jan 1974 
16 3 389-610 28 Aug 1995 2 : pon )eo: SN IAS 
16 2 225-388  30Dec 1994 5 ie A OMal Ia? 
16 1-222 26 Aug 1994 4 4 279-430 28 Sep 1972 
15 4 565-669 29 Dec 1993 4 3 228-278 = 22 Jul 1971 
15 3 349-563 28 Sep 1993 ‘i 2 ae EBL 
15 2 173-348 31 Dec 1992 2 I-36 16 Sep 1968 
15 1 1-172 17 Sep 1992 3 7 451-444 3 Apr 1970 
14 4 525-638 31 Dec 1991 : ° ead dade 
14 3 315-524 27 Aug 199] 2 a 
14 2 141-314 27 Dec 1990 3 : 191-283 21 Jun 1968 
a ae. oes 3 3 123-190 24 Nov 1967 
3 2 77-122 31 May 1967 
13 4 393-518 29 Dec 1989 3 1 _76 31 May 1967 
13 3 265-392 31 Aug 1989 
13 2 125-264 30 Dec 1988 7 6 are =O Der Tee 
ie oe 2 5 349-394 27 Sep 1966 
2 4 269-348 20 Jul 1966 
12 4 Never published 2 3 177-268 Apr 1966 
12 3 Never published 2 2 115-175. 31 Dec 1965 
12 2 253-434 30 Dec 1987 2 1 1-113 3 Mar 1965 
12 1 1-251 29 May 1987 
1 é 299-417 30 Oct 1964 
11 4 357-497 16 Dec 1986 1 5 957-298 25 Jun 1964 
11 3 2507250 20 May 1986 1 4 187-256 25 Jun 1964 
11 2 107-253 12 Dec 1985 1 3 109-186 12 Dec 1963 
11 ] 1-106 19 Jun 1985 1 2 63-108 23 Nov 1962 
1 1 62. 23 Nov 1962 


A NEW SPECIES OF WITCH-HAZEL 
(HAMAMELIS: HAMAMELIDACEAE) 
APPARENTLY ENDEMIC TO SOUTHERN MISSISSIPPI 


S.W. Leonard 


The Nature Conservancy 
Camp Shelby Field Office 
CSTS-ENV, Bldg. 6678 
Camp Shelby, Mississippi 39407, U.S.A. 


ABSTRACT 


A new species of witch-hazel, Hamamelis ovalis, is described from Perry County in southern Mississippi, US.A 
The red flower color, clonal habit, and late: winter prOMeNng Sugeest affinity with the Ozarkan endemic H. ve ridilis 
however, the large leaves, unlike a merican or A witch-hazels, readily differenti- 


a 


ate H. ovalis from its congeners. The problematic H. macrophylla is discussed. 


RESUMEN 


Se describe una nueva especie de la avellana-bruja, Hamamelis ovalis, del condado de Perry, Mississippi del sur, 
los Estados Unidos. El color rojo de la flor, el habito conics. y ie norcen radi en i g aunadad 


con el Ozarkan endémico H. vernalis. Sin embargo, | 
de avellanas-brujas americanas 0 asiaticas eine Poiimente H. ovalis de sus Weoneencies Se discute el 
problematico H. macrophylla. 


During surveys for proposed tracked vehicle maneuver areas at Camp Shelby Training Site 
in southern Mississippi, an unusual colony of witch-hazel (Hamamelis) was discovered in 
July 2004. Distinctive features initially observed were the large, abaxially pale whitish- 
green ovate leaves, densely pubescent cinnamon-brown twigs, and abundance of low clonal 
sprouts. The colony was periodically checked to see when anthesis would occur and if the 
petals would be yellow like all the known native Hamamelis plants in southern Missis- 
sippi. In early January 2005 the large tawny buds opened to expose brilliant red sepals and 
petals entirely maroon, scarlet, rose, or red with yellow tips. Subsequent surveys in the area 
revealed colonies with mixed flower colors as well as colonies with flowers entirely wine- 
red. Leaf and floral appearance as well as the clone-forming habit of the plants did not 
match anything known for Hamamelis from the Gulf Coast region. 

Witch-hazel shrubs, with pale yellow petals less than 8 mm long and occasional 
plants with leaves less than 6 cm long are present at Camp Shelby. These yellow-flowered 
plants do not fit this writer’s concept of Hamamelis virginiana L. var. virginiana of the 
Carolinas nor Hamamelis virginiana L. var. henryae Jenne ex C. Lane of northern Florida. 
However, for simplicity of discussion in this paper, they will be referred to as H. virginiana. 
Floristic treatments indicate a range for H. virginiana from Nova Scotia to Texas (Small 
1933; Fernald 1950; Steyermark 1963; Radford et al. 1968; Godfrey 1988; Meyer 1997) and 
possibly Mexico Jenne 1966; Lane 2005). In Mississippi these shrubs and small under- 
story trees are frequent on acidic, sandy woodland slopes and bluffs, moist or dry, in full 
sun or shade. Flowering at Camp Shelby begins in November, peaks in December and tapers 
off in January. In addition to the native woodland Hamamelis virginiana in Mississippi, 
other witch-hazels are occasionally used as landscaping shrubs. These cultivars include 


SIDA 22(2): 849 — 856. 2006 


850 BRIT.ORG/SIDA 22(2) 


the Ozarkan witch-hazel, Hamamelis vernalis Sargent, and less frequently the Oriental 
species, Hamamelis mollis Oliver, Hamamelis japonica Siebold and Zuccarini, and the 
hybrid Hamamelis x intermedia Rehder. Neither H. vernalis, nor the Asian plants are 
known to naturalize in Mississippi. 

In March 2005 the dried collection of Hamamelis at the Missouri Botanical Garden 
Herbarium (MO) and late flowering specimens of H. vernalis Sargent on the Garden’s 
grounds were examined. Although considerable variation in foliage was noted, nothing 
in the MO collection approximated the big-leaved, red-flowered plants found earlier at 
Camp Shelby, Mississippi. With no visible evidence that the Camp Shelby colony was 
located near a house site where cultivars might have been introduced, exotic witch-ha- 
zels were eliminated. Leaf size, pubescence, clonal habit, flower color, petal length, and 
phenology tended to rule out the common eastern witch-hazel, H. virginiana—highly 
variable as it is known to be. 

Clonal habit and infusion of red in the petals suggested an affinity between the Camp 
Shelby plants and the variable Hamamelis vernalis,an Ozarkan endemic that grows 600 
kilometers to the northwest of the Mississippi site. Rehder (1920, 1928) and Steyermark 
(1963) recognized forms of H. vernalisto account for the pure red flower color and tomen- 
tose leaves within certain colonies of H. vernalis. Jenne (1966) proposed that the variable- 
flowered, clonal plants of H. vernalis originated as putative hybrids between the eastern 
H. virginiana and an unknown parental type (possibly no longer extant). However, 
Bradford and Marsh (1977), af ter ] parative studies in Arkansas of H. virginiana 
and H. vernalis, were inclined to disagree with Jenne’s interpretation of the hybrid origin 
of H. vernalis. Nevertheless, as Jenne (1966) hypothesized, a tentative Ozark relict “would 
be expected to be extreme in the following directions: short petal length, presence of red 
pigment in the petals and on the inner surface of the sepals, linear staminodia, less than 
0.5 mm wide, the tips of which are not dilated, clonal habit of vegetative suckering, with 
75 to 100 stems emerging from one root system.” Looking at these features only, the Camp 
Shelby plants come uncannily close to Jenne’s parental phenotype, but size and vestiture 
of leaves, branching pattern, habitat, and other characteristics are hard to reconcile the 
Camp Shelby shrub asa parent of H. vernalis. 

On the other hand, with exception of variation in petal color, nothing about the Camp 
Shelby plants suggests a hybrid origin. In fact, other features such as larger leaf size, ear- 
lier flowering, musty floral fragrance, plant habit, and dry pineland habitat point to- 
ward an independently derived species. As documented below the Mississippi plants are 
sufficiently different from the Ozark plants to merit a taxonomic designation at specific 
rank. Therefore the following name and diagnosis are provided: 


ee ovalis S.W. Leonard, sp. nov. (Figs. 1-2). Type: U.S.A. Mississippr Perry Co. shallow 
vine in mixed pine-hardwoods near headwaters of Garraway Creek, Camp Shelby Training 
— T-17, Compartment 96, DeSoto National Forest, T2N, R1W, Sec. 6, NEI/4, 7 Jul 2004, 

S.W. Leonard 11073 (HOLOTYPE: MO, ISOTYPES: FSU, NCU, NY). 


Frutex clonis, crescit ad 5 m. Differt a Hamamelis virgini ) quod scarlatinus vel rubropurpureus est, 


floris folia sunt coloris vinosi aut rosei; differt a H. ver nelas e0 ty folia magna ovata habet et pars obversa folii 
habet pilos stellatos albos. 

Rhizomatous shrubs to 5 m tall with twigs densely stellate pubescent, brown to brown- 
ish-gray on older twigs; terminal buds russet. Leaves tardily deciduous, alternate, ovate 
12-24 cm long, 5-17 cm wide, dark green and scurfy above, pale green with dense stellate 
tomentum below initially white and aging to golden tan, margins sinuate on distal half 


LEONARD, A NEW SPFEAICS UF 851 


Fig. 1. Powel vet) eee ovalis, ii, Cee ken on January: 2006.1 (right) j | yp 
ality eruleris 15 c I } jal £ dq + | hy bnit 


J J J 


and entire on proximal half; petioles 1.0-3.5 cm long; leaf bases oblique with longer lobe 
rounded to cordate, principal veins of leaves alternate, usually 11 (five on one side of mid- 
rib, six on the other); leaf apices acute to obtuse. Flowering late December to early Febru- 
ary with peak around January 20; calyx of 4 ovate sepals 3-4 mm long, acute, densely 
tawny pubescent abaxially and glossy scarlet adaxially, spreading, slightly reflexed, and 
ultimately strongly reflexed with age, subtended by 2 or 3 densely tawny tomentose 
bractlets; petals liguliform, at first uncoiling 7 mm long and continuing to lengthen to 14 
mm over a period of 3 weeks, circinnately coiled in bud and irregularly revolute when 
expanded, various shades of red (wine-red, scarlet, rose) or infrequently tipped with yel- 
low; filaments purplish-red, gradually expanded toward apex and resembling gavels until 
anthers turn outward from the connective exposing sockets in the distal ends of the fila- 
ments; filaments alternating with staminodes; staminodes dilated distally and curved 
outward with the broader tips straddling the base of each petal, thus pushing the petal 
away from the stigmatic surface; ovary partly inferior, pubescent with mostly appressed 
hairs, fruits slightly compressed ca. 2 cm long calyx adhering to capsule except for re- 
curved tips; seeds 2 per locule, glossy black, 7-10 mm long, slightly obovate with each 
face of attachment scar (2 mm wide and 3 mm long) about twice the width and length of 
attachment scars on seeds of H. virginiana from local populations (Fig. 1). 

Ecology and phenology.—At the type locality, the plants occur in a shallow flat-bot- 
tom ravine and on adjacent low slopes. Soil at the site is loamy. The ravine drains into a 
headwater tributary of Garraway Creek that flows into the Leaf River between the City 
of Hattiesburg and the small municipality of New Augusta in Perry County, Mississippi. 
The location is in Compartment 96 of DeSoto National Forest, Training Area 17 of Camp 
Shelby Training Site, T2N, R11 W, Section 6. 

The Hamamelis occurs beneath a canopy of pine and mixed deciduous hardwoods. 
The main species of this vegetation are: Pinus taeda, Pinus palustris, Carya tomentosa, 
Nyssa sylvatica, Quercus falcata, Magnolia macrophylla, Cornus florida, Sassafras albidum, 
Ilex vomitoria, Ilex opaca, Liquidambar styraciflua, Vaccinium arboreum, Vaccinium 
elliottii, Morella cerifera, Callicarpa americana, Tridens flavus var. chapmanii, 


852 BRIT.ORG/SIDA 22(2) 


Fic? Clockwi £ ra Late. U iB ae a (F fa MS). H yi Th (P rf. MS) H, Tj, Th. (T ‘a 
. L 


MO), Hamamelis virginiana (Perry Co., MS). Each 


LEONARD 853 


Chasmanthium sessiliflorum, Sorghastrum elliottii, Smilax pumila, Mitchella repens, 
Berchemia scandens, and Vitis rotundifolia. 

Flowering of Hamamelis ovalis was observed in late December with the peak occur- 
ring around the first ten days of January in 2005. The 2006 peak (during an unusually 
warm January) was closer to January 20th. In addition to the taller plants developing 
flowers, clonal sprouts—some no more than 20 cm tall—also flower. 

The following observations were made in 2006 on potted plants of H. ovalis and H. 
vernalis: flowering of H. ovalis begins (late December) as flowering of H. virginiana is 
ending; flowering of H. vernalis begins (mid-February) as flowering of H. ovalis is end- 
ing. Leaves of H. ovalis are 30-40 percent of mature size before the leaves of H. vernalis 
begin to emerge from the buds. 

Habit and morphology.—Although Hamamelis ovalis, like H. vernalis, isa clonal shrub, 
the root sprouts are generally uniform in height on upper slopes where prescribed fire at 
2-6 year intervals isa standard longleaf pine management procedure. These small shoots, 
typically 3-5 dm tall, extend outward as muchas 6-8 meters from the central plant. How- 
ever, the habit of the H. ovalis is markedly different from that of H. vernalis. Instead of 
forming a rounded bush of several sucker stems with major branches diverging in all 
directions from near the base of the plant as in H. vernalis, mature shrubs of H. ovalis 
have two growth forms—distichous branching ona geniculate central axis and a graceful 
arched branching on an erect straight stem. On the former, alternate branches diverge at 
approximately right angles in the same plane, gradually becoming longer as they progress 
up the main stem, thus giving the plants an espaliered look. Rhizomes of H. ovalis are 6- 
10 mm in diameter and are shallow, often just beneath the leaf duff. In contrast, rhizomes 
of H. vernalis are deep seated among overwash stones in flood zone sands. Specimens 
along the Caddo River in Clark County, Arkansas were tightly bound to the substrate 
and not easily extracted. 

The type locality colony of Hamamelis ovalis comprises 92 stems greater than 1.5m 
tall with the tallest being 3.8 m (%2.3 m) and about 120 smaller sprouts. It is estimated 
these 200+ stems arise from the rhizomes of fewer than two dozen plants. Field work 
during 2005 in the headwaters of Garraway Creek revealed about two dozen smaller colo- 
nies of witch-hazel in drier habitats: upper slopes, crests of ridges, and 1-2 hectare level 
areas now covered with mixed longleaf and loblolly pine secondary growth forests. 

Calyces are consistently deep reddish-maroon adaxially with conspicuous tawny 
hairs from the dense abaxial pubescence protruding along the lobe margins. Calyces of 
some plants become tinged with green toward the base of the lobes after the petals wither. 
Petals were predominantly wine-red, scarlet, and reddish-pink, with at least one plant 
having yellow petals with the basal half deep red. Petals persist for around three weeks 
before they begin to turn brown from the tips progressively toward the base. Floral fra- 
grance of H. ovalisis at first faintly spicy or “fruity,” thereafter degrades to a musty scent 
similar to that of decaying pears, and eventually to a disagreeable “wet dog” odor! 

One of the distinctive features of Hamamelis ovalis is the abaxial appearance of ma- 
ture leaves in the early part of the growing season. If viewed at an angle in direct sun- 
light, the developing leaves have a hoary white appearance above due to closely spaced 
stellate hairs. At the same time the lower surface is whitish (very pale green) from closely- 
knit stellate pubescence. 

On plants of Hamamelis ovalis, even short sucker sprouts, two or three of the leaves 


854 BRIT.ORG/SIDA 22(2) 


toward the stem tip will be greater than 20cm in length. This dimension exceeds maxima 
for all witch-hazel taxa currently described. In autumn the leaves of H.ovalis turn yellow, 
then brown, and many continue to hang downward on the branches until mid-March. 
Anobservation that may have some bearing on data collection from herbarium speci- 
mens is that witch-hazels respond vigorously after forest fires. At Camp Shelby Training 
Site, stems of Hamamelis virginiana in the first one or two growing seasons after fire 
have larger than normal leaves, usually 16-18 cm long. These post-fire leaves tend to have 
cordate rather than oblique leaf bases and lack conspicuous abaxial pubescence. 


Hamamelis ovalis vs. H. vernalis 


. Leaves ovate, 12-24 (%19.7) cm long, 5-17 cm wide, soft velvety green above for first half of 
growing season and ae becoming darker green ae SC ay above; pale green va ae 
dense white tomentum in spring that graduall nin autumn, 
sinuate on distal half, base rounded, strongly cordate-oblique, usually 11 major veins arising 
from midrib with 5 on one side and 6 on the other; suckers from base of main stem absent; 
plants strongly soboliferous with rhizome sprouts mostly less than 5 dm tall; flowering late 
December to early February, flowers faintly spicy becoming musty for the three-week duration 


ez 


of the petals H. ovalis 
. Leaves mostly obovate, 7-13 (%=8.7) cm long and 4-10 cm wide glabrous above or sparingly 

stellate pubescent becoming darker green and smooth above; ce ous beneath or rusty stel- 

late pubescent along midrib and veins, margins sinuate apically, cuneate basally or infrequently 

weakly rounded and oblique, usually 9 or 10 major veins arising nae midrib; plants suckering 

strongly from base and occasionally sending up rhizome sprouts that form dense thickets 2-3 

meters tall; flowering late February throughout March, flowers sweetly aromatic H. vernalis 


Leaves of Hamamelis ovalis are more than twice as large as leaves of H. vernalis, and sel- 
dom if ever have an obovate shape. In contrast, foliage of H. vernalis observed along the 
Caddo River in Arkansas as well as leaves of cultivated plants of this species are consis- 
tently obovate, slightly glaucous, and with broad cuneate bases. Flowers of Hamamelis 
ovalis are similar in size and possibly as variable in color as H. vernalis, but lack the pro- 
nounced fragrance. 
Hamamelis ovalis vs. H. virginiana 
arge ovate leaves with pale densely stellate tomentum on abaxial surface; mostly 11 lateral 
veins per leaf; abundant short rhizome sprouts sometimes forming a groundcover; tawny pu- 
bescent twigs of current and previous season's pe flower buds and terminal buds larger, 
scarlet adaxial surface of calyx; maroon, scarlet, or deep pink petals and larger; flowering princi- 
pally in January in Mississippi; meee nat seeds with larger attachment scars H. ovalis 
. Leaves 3.7-16.7 cm long and 2.5-13 cm wide (Meyer 1997); elliptic, obovate, nearly rounded; 
margins sinuate, base broadly cuneate, ned weakly to strongly oblique with one lobe 
rounded the other cuneate; surface dull green above, scabridulous or smooth, lower nae 
pale green, glabrous or pubescent or tomentum restricted to midrib and major veins; 9 or | 
ateral veins arising from midrib; plants suckering from base with only a few sprouts arising 
from rhizomes; flowers pale yellow, smaller (6-8 mm) than published dimensions [10-20 mm 
see Meyer 1997)] for the species, opening in November to mid-December 1s slight srl 
with smaller attachment scars H. virginiana 


1. 


cz 


Features that differentiate Hamamelis ovalis from the widespread eastern North American 
H. virginiana are the larger leaves with pale densely stellate tomentum on abaxial surface; 
tawny pubescent twigs, longer bud scales, scarlet adaxial surfaces of calyx, reddish petals 
instead of typical yellow, floral fragrance, and strong development of clonal sprouts. Flow- 
ering of H. ovalis begins ate December) as flowering of H. virginiana is ending. 

Among the nearly two dozen discrete colonies of Hamamelisovalis observed at Camp 


LEONARD 855 


Shelby, only one has been found that contains both H. ovalis and H. virginiana (small leaf 
form) and at this site there are only a few plants of the latter. In areas where Hamamelis 
ovalis occupies drier sites at the heads of drainages, H. virginiana tends to be a stream- 
side plant. 


Identity of Hamamelis macrophylla 

Because the large leaf size of the new Mississippi witch-hazel is one of its most distinc- 
tive features, a logical name for the shrub would be Hamamelis macrophylla. This name 
was in fact used by John Lyon, a Scottish nurseryman working out of Philadelphia and 
making commercial collecting trips as far south as Georgia and Florida. Where Lyon found 
his new species is unknown. He offered it for sale in 1812 under the name Hamamelis 
macrophyllus sp. nova, but he did not publish a Latin diagnosis (Ewan and Ewan 1963). 
Pursh appropriated the name, provided a brief Latin diagnosis, and published the name 
in his Flora americae septentrionalis (Pursh 1814). Although Pursh said the type came 
from “river banks in the western part of Georgia,” Ewan and Ewan (1963) pointed out in 
a footnote to the annotated Lyon journal that some of Pursh’s locations were not “trust- 
worthy.” Loudon, a Lyon biographer writing in 1844, according to Ewan and Ewan (1963), 
said the witch-hazel was native to the “Katawba (sic) Mountains.” This location would 
place the type locality in western North Carolina. 

Regarding the Lyon/Pursh Hamamelis macrophylla, Pursh described only “large 
leaves punctated on their lower side, with rough tubercles and other marks...” Appar- 
ently he never saw flowers of H. macrophylla and if Lyon saw the plants in flower, he 
either withheld the information from Pursh or omitted it from his journal of 1799-1814. 
Pressed specimens were either lost or destroyed. 

A clue regarding the provenance of Hamamelis macrophylla might be ascertained 
from associated flora. Having seen the magnificent Magnolia macrophylla, Lyon seemed 
to have become obsessed with it and is said to have collected more than 4,000 seedlings 
from a single location in North Carolina (South Carolina Association of Naturalists news- 
letter June 1996). It is possible that on one of his collecting trips to Magnolia colonies in 
western Georgia (or western North Carolina?) he collected a large-leaved Hamamelis. 
According to Lyon’s notes of his Georgia travels, he was near the Ocmulgee River on June 
23,1803, the Flint River on July 17, 1803, and the Chattahoochee River on October 6, 1803. 
However, these dates appear to be too early if the witch-hazels were not advertised for 
sale until 1812. A more likely site and date of collection would be western North Carolina 
near the Catawba River in October 1809, when Lyon packed 3,600 seedlings of Magnolia 
macrophylla tor shipment to Europe. 

agnolia macrophylla is a frequent associate of Hamamelis ovalis in the upper 
Garraway Creek watershed in Mississippi. If the Georgia or the Carolina plants bloomed 
in late winter or spring, were red-flowered and were clonal, then Pursh’s Hamamelis 
macrophylla might be the same species as recognized here as newly described. If the Mis- 
sissippi plants were a colony of a conjectured red-flowered H. macrophylla, such a dis- 
junction would be about 425 kilometers from a theoretical site in Georgia or more than 
1100 km from a North Carolina location, based on the distribution of Magnolia 
macrophylla and assuming the magnolia’s range has changed little since the early nine- 
teenth century. 

Treatments of the common eastern Hamamelis virginiana have included Hamame- 
lismacrophylla Pursh as asynonym (e.g, Sargent 1922; Coker & Totten 1945). Unless, how- 
ever, John Lyon’s steps can be retraced to an extant colony of a big leaf witch-hazel, or 


856 BRIT.ORG/SIDA 22 


— 


2) 


unless nursery stock from Lyon’s 1812 sale has been propagated for almost two centuries 
in an English garden, the true identity of H. macrophylla may never be known. 


ACKNOWLEDGMENTS 


My thanks to Alan S. Weakley, NCU Herbarium Curator, for searching references tucked 
away in the University of North Carolina archives and to Mary Stevens of the Mississippi 
Museum of Natural Science for obtaining the paper by Bradford and Marsh. Lisa Yager, 
Camp Shelby Field Office Coordinator, arranged interlibrary loans of two crucial docu- 
ments. Suggestions by reviewers, Guy Nesom, Bruce Sorrie, and Stephen Spongberg greatly 
improved the manuscript. | thank Fr. Peter Verhalen for corrections to the Latin diagno- 
sis, the Mississippi Army National Guard for funding, and the US Forest Service, DeSoto 
Ranger District, that has final management responsibility for conservation of the species 
and its habitat. 


REFERENCES 

Braprord, J.L.and D.L. Marsu. 1977.Comparative studies of the witch hazels Hamamelis virginiana and 
H. vernalis. Proc. Arkansas Acad. Sci. 31:29-31. 

Coker, W.C.and H.R. Torten. 1945. Trees of the southeastern states. Univ. of North Carolina Press, Chapel 
Hill, North Carolina. Pp. 195-196. 

Ernst, WR. 1963.The genera of Hamamelidaceae and Platanaceae in the southeastern United States. 
J.Arnold Arbor. 44:193-210. 

Ewan, J.and N. Ewan. 1963. John Lyon, nurseryman and plant hunter, and his journal, 1799-1814. Trans. 
Amer. Philosophical Soc. Vol. 3, pt. 2. Philadelphia, PA. 

Fernato, M.L. 1950. Gray’s manual of botany. American Book Company, New York. P.752. 

Goorrcy, R.K. 1988. Trees, shrubs and woody vines of northern Florida, and adjacent Georgia and 
Alabama. Univ. of Georgia Press, Athens. Pp. 372-374. 

Jenne, G.E. 1966. A study of variation in North American Hamamelis L. (Hamamelidaceae). Master of 
Arts thesis, Vanderbilt Univ., Nashville, TN. 

Lane, C.2005.Witch hazels. Timber Press, Inc., Portland, OR. 

Mever, E.G. 1997. Hamamelidaceae. In: Flora of North American Editorial Committee, eds. Fl. North 
Amer. Oxford Univ. Press, New York and Oxford. 3:362-36/7. 

Pursu, FT. 1814. Flora americae septentrionalis. London,White, Cochran and Co,, London, Great Britain. 
1:116. 

Raprord, A.E.,H.E. Antes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. Univ.of North 
Carolina Press, Chapel Hill. Pp. 529-531. 

Revper, A. 1920. New species, varieties, and combinations. J. Arnold Arbor. 1:256. 

ReHper, A. 1928. New species, varieties, and combinations. J.Arnold Arbor. 9:30. 

SarGENT, C.S. 1922. Manual of the trees of North Americ a. Houghton, Mifflin and Co., Cambridge, MA. 
1:368-371. 

Smatt, J.K. 1933. Manual of the southeastern flora (1972 facsimile of the 1933 edition). Hafner Pub- 
lishing Company, New York. Pp. 600-601. 

South CAROLINA ASSOCIATION OF NATURALISTS. 1996. June newsletter. Columbia, South Carolina. 

SreverMaRrK, JA. 1963. Flora of Missouri. lowa State Univ. Press, Ames, lowa. Pp. 786-789. 


ERIOGONUM CALLISTUM (POLYGONACEAE), A NEW SPECIES 
FROM THE TEHACHAPI MOUNTAINS OF CALIFORNIA 


James L. Reveal! 


ersity of Maryland 
College ee a and 20742-5815, U.S.A. 
and 
New } rk Botanic Cd! rde! 
Bronx Park, New York 10458- 5026 USA. 
jreveal@umd.edu 


ABSTRACT 
Eriogonum callistum, a new species of subg. Eucycla Nutt. from the Tehachapi Mountains of Kern Co., California, 
is the sole representative of Sect. Lanocephala, a new section allied to Sect. Lachnogyna Torr. & A. Gray, a taxon 
pans al ne acre ce but probably more osely revatee to Sect Pal sonia oy ot the agile peak The 


gla rg eC compact ensel 
ae new pieaves from which arise several UpHent poapoee flowering s stems terminated by a capitate inflorescence 
subtended by many bracts with many g hairy, bright white flowers on long, hairy 


pedicels, and glabrous, trigonous achenes. 


RESUMEN 


Eriogonum callistum, una nueva especie del subg. Eucycla Nutt. ede las montanas Tehachapi de Kern Co., Califor- 
nia, es el Unico pep ieschiante oS la Sect. Panocephala, | ite Sect. Lachnogyna Torr. & A. 
babl 4 a Sect Lattjola 


Benth. de la costa pacifica. La nueva especie ee port formar grand i lond 


hojosas y rosetas de hojas nuevas, de las que I iba: llos florid 
una inflore ia capitada subtendida por muchas bracteas, con muchos involucros ae een numerosas 


flores blanco brillante, sobre pedicelos largos y pelosos, y aquenios trigonos glabros. 


The discovery of another undescribed member of Eriogonum Michx. (Polygonaceae Juss., 
subf. Eriogonoideae Arn.) from California is not, in itself, a surprise, but one so distinct as 


to represent a new section, and of such potential as a cultivated garden plant, is at least 
unusual. 


Eriogonum Michx. Sect. Lanocephala Reveal, sect. nov. Type: Eriogonum callistum Reveal. 

A Eriogono sectionibus singularis floribus et pedicellis pilosis. 

sa ea callistum Reveal, sp. nov. (Fig. 1). Typr:U.S.A.CALIFORNIA: Kern Co: Tehachapi Mountains, 
y 2006, R.E. Preston 2400 (HOLOTYPE: UC; IsoryPes: NY, RSA, U 

A Eriogono lachnogyno Torr. ex Benth. foliis latioribus et longioribus, involucris numerosis (10-35 nec 2-5), et 

floribus albis (nec flavis) et E. latifolio Sm. floribus pilosis differt 

Plants perennial forming densely white-tomentose, rounded, compact mounds, 3-10(-11) 

dm across and (0.5-)1-3.5 dm high, arising from a stout, woody taproot with numerous, 

spreading, tightly compact caudex branches covered with marcescent leaves ultimately 

terminated by a tight leafy rosette of numerous, newly-formed basal leaves; leaves fas- 

ciculate in terminal tufts, persistent, the petiole 1.5-5 cm long, grayish-white tomentose, 

the petiole-base narrowly triangular, glabrous, 0.5-3 cm long, 0.3-0.8 cm wide, becoming 


'Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, U.S.A. 
SIDA 22(2): 857 — 861. 2006 


858 BRIT.ORG/SIDA 22(2) 


a7 
G 
§ 
\ 
» 


{ 


yy 

a 
\ 

‘A 


!y 
y 


x 


\ 


We 


oe a wy 
Mbsuwey 


Fic. 1. Illustration (lef ight) of a leaf, an involucre, and a flower with its pedicel and bractlets. 
~s Pp 


papery and reddish-brown with age, the blade elliptic, flattened or slightly folded longi- 
tudinally, (1-)2-5 cm long, 0.8-2 cm wide, silky grayish-white tomentose on both sur- 
faces, the margins entire, flat; flowering stem scapose, arising centrally from rosette, erect 
or nearly so, slender, solid, not fistulose, (1.5-)2-4(-4.5) dm long, greenish to grayish, be- 
coming reddish with age, thinly tomentose; inflorescences capitate, 2-4 cm across, bracts 
8-16, subfoliaceous, linear to narrowly lanceolate, 4-12(-15) mm long, 1-3 mm wide, to- 
mentose on both surfaces except glabrous basally on adaxial surface; peduncles absent, 
involucres 10-25(-35) per cluster, turbinate, (5-)6-8(-9) mm long, 2-4 mm wide, rigid, 
tomentose abaxially, glabrous adaxially, the teeth 5, erect, 2-3.5 mm long, acute apically; 
flowers bisexual, not attenuated basally, 2-5 mm long, the pedicels pilose, (5-)6-10(-12) 
mm long, the bractlets linear, 4-8(-10) mm long, pilose throughout, the perianth pink- 
ish-white in bud, becoming bright white, densely white tomentose with long, soft hairs 
abaxially, glabrous adaxially except for glands along the rosy to yellowish-green mid- 
ribs, the tepals connate |/4-1/3 their length, essentially monomorphic, oblong to ellip- 
tic, the outer three slightly broader than the inner three; stamens exserted, 3-6 mm long, 
the filaments pilose proximally, the anthers oblong, pale yellow with a tinge of pink to 
light rose, 0.4-0.5 mm long; achenes trigonous, 3-4 mm long, glabrous. 


Other specimens examined: U.S.A. Kern Co.: Tehachapi Mountains, 13 Jun 2006, B.D. Schafer 214 (DAV, NY, RSA, 
UC, US); 25 Jul 2006, [-L. Reveal et al. 8781 (NY, RSA, UC, US). 


~ 


Eriogonum callistum (from kalliston Gr. most beautiful; Figs. 2, 3) represents a new sec- 
tion within subg. Eucycla Nutt. The Sect. Lanocephala (from lenos Gr. wooly hairs, and 


REVEAL, A NEW SPECIES OF ERIOGONUM FROM CALIFORNIA 859 


cae 
srbete tae 


A ae 
“Hee 
> 


860 BRIT.ORG/SIDA 22(2) 


kephale, head, as to the pilose flowers on hairy pedicels arranged in a capitate inflores- 
cence) is similar to Sect. Lachnogyna Torr. & A. Gray, a taxon of two species encountered 
from eastern Arizona to western Texas northward onto the southern Great Plains to Colo- 
rado and Kansas. Although the type, E. lachnogynum, has hairy achenes, the other spe- 
cies, E. havardii S. Watson, has glabrous fruits. The soft, silky, grayish-white tomentum 
of the leaves of E. callistum is similar to that seen in both species, and while both mem- 
bers of Sect. Lachnogyna have yellow flowers, those of E. callistum are a bright white. The 
monospecific section is probably more closely allied to Sect. Latifolia Benth., and specifi- 
cally the coastal element of the taxon, E. latifolium Sm. Both form rounded mounds of 
leaves and have numerous involucres in capitate heads. Still, E. latifolium is best charac- 
terized asa subshrub or shrub with long, aboveground, woody stems. Eriogonumcallistum 
is an herbaceous perennial with short, stout, caudex branches buried in a mixture of soil 
and marcescent leaves. Nonetheless, Sect. Lanoce phala is the only taxon in the genus with 
hairy pedicels. The flowers of E. callistum are individually small but numerous in nu- 
merous involucres that together form a capitate cluster atop long, slender scapes. The 
achene is trigonous, glabrous, and contains a curved embryo in abundant mealy en- 
dosperm; as such it is typical of most species of subg. Eucycla. Ongoing molecular stud- 
ies should resolve an exact placement of Sect. Lanocephala. 

At present, Eriogonum callistum is known only from a few scattered locations that 
altogether contain some 2000 individuals in a variety of age classes. The new species 
grows mainly in open places among manzanita on rock outcrops of pre-Cretaceous lime- 
stone (Jennings & Strand 1969), on open ridges within mixed chaparral. Associated spe- 
cies include Adenostoma fasciculatum Hook. & Arn., Amelanchier utahensis Koehne, 
Arctostaphylos glauca Lindl, Arctostaphylos parryana Lemmon, Hesperoyucca whipplei 
(Torr) Trel., Pinus monophylla Torr. & Frém., Ceanothus cuneatus (Hook.) Nutt., Quercus 
berberidifolia Liebm., Tauschia parishii J.M. Coult. & Rose) J.F Macbr, Castilleja martinii 
Abrams, and Corydanthus eremicus (Coville & C. V. Morton) Munz. 

In the key to Eriogonumin Flora of North America (Reveal 2005), the new wild buck- 
wheat will key to subg. Eucycla, Key 1-California (p. 225) and will proceed to lead num- 
ber 12 (p. 226). That may be modified as follows: 

12. Perianth glabrous; s Sierra Nevada, Transverse Ranges, and desert ranges to the east from 
ono County s to Los Angeles and San Bernardino counties 73.E pias kennedyi 
12. Perianth pubescent; Last Change Range, Inyo and Mono counties, or Tehachapi coe 
Kern County. 
12a. Achenes pubescent; pedicels glabrous; scapes up to 0.3 dm long; Last Change Range, 
Inyo and Mono counties 68.E dee shockleyi 
12a. oe glabrous; pedicels pilose; scapes 2—3.5 dm long; Tehachapi wen Ker 
County 92a 


— 


Mm Ilictiim 


The author has agreed not to publish at this time detailed information as to the exact 
location and distribution of Eriogonum callistum. 

Eriogonum callistum should make an attractive addition to the rock garden flora. 
The large, roundish mats are composed of numerous tight rosettes of persistent leaves 
that remain attractive long after flowering. The fresh leaves, plus those of previous years 
that remain on the caudex branches, allow the mats to accumulate soil and plant debris 
and thus enlarge as roundish mounds with each growing season. The numerous, closely 
arranged caudex branches mean that the plants can be grown as tight, compact clusters 
along borders or in patterns. The plant’s natural habitat suggests that it requires little 


REVEAL, A NEW SPECIES OF ERIOGONUM FROM CALIFORNIA 861 
watering, prefers gravelly soil in open, sunny places, and should survive with little or no 
maintenance. 
ACKNOWLEDGMENTS 
I wish to thank Hazel Pollard for preparing the line drawing. 
REFERENCES 


JENNINGS, C.W. and R.G. STRAND. 1969. Geologic map of California, Los Angeles Sheet. California Division 
Mines and Geology, Sacramento. Scale: 1:250,000. 

Reveal, J.L. 2005. 44a. Polygonaceae Jussieu subfam. Eriogonoideae Arnott, Encycl. Brittannica (ed. 7), 
5:126. 1832. Wild buckwheat. Fl.N. Amer. 5:218-478. 


BRIT.ORG/SIDA 22(2) 


Book REVIEW 


Guy GusMAN and LILLIAN GusMAN. 2006, The Genus Arisaema, a Monograph for Botanists 
and Nature Lovers, Second Revised and Enlarged Edition. (ISBN 3-906166-37-6. Publisher: 
ARG Gantner Verlag Kommaditgellschaft FL 9491 Ruggell. Distributor: Koeltz Sci- 
entific Books, POB 1360, D-61453 Konigstein/Germany. (Orders: Timber Press Inc, 
133S.W Second Avenue, Suite +50, Portland, OR 97204-3527, U S.A; wwwtimber press.com, 
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). US$79.95 
and C$109.95. +74 pp., 400 +color ee 29 figures, and 18 tables, 6 3/8" x 91/2". 


ribed since ene First Edition was published, many 


This Second Edition includes more than ten new s] 
new photographs, geographical distribution of species by country, d 
Sevelopaicatid in Arisaema taxonomy, types of acai taxa 4 ie d piles aphical references. 

‘The genus Arisaema consists of about 200 species, subspecies and varieties whose extraordinarily wide 
range extends from Central and East Africa towards Southeast Asia, via Yemen, Oman, Pakistan, Afghanistan, 
the whole Himalayan range, India, China, Korea, Japan and Siberia and auras to North America. Arisaema 
is the third largest of the 105 genera that make up the Araceae (Mayo, Bogner & Boyce), 1997), a remarkable fam- 

ily of monocotyledons that contains some of the most extraordinary ee in the world.” (pg. 12) 

This, a second, revised and enlarged edition, is a comprehensive book on the genus, written by two collec- 
tors and growers, who are themselves scientists. Commonly called Cobra-Liles, most do not resemble a snake 
and the genus is unrelated to lilies. The initial chapters include ees ee use in chemistry and medicine, 
occurrence in the wild and in cultivation, Beas ation and systematics. Most of the book is taken up in the de- 
ections, which includes photographs, drawings, tables, etc. Chapters on 


tables for key species, recent 


scription of species by each of the 
doubtful species, invalid names and then hy ae follows. The final three chapters include a ee bibliogra- 
phy, and an index to species names. 

The book is a must have for botanists, horticulturalists, plant collectors, ee growers, and anyone 


interested in the fascinating genus, Arisaema.—David Leedy, Volunteer, Botanical Research Institute of Texas, Fort 


Worth, TX 76102-4060, U.S.A. 


SIDA 22(2): 862. 2006 


SIX NEW COMBINATIONS IN DODECATHEON (PRIMULACEAE) 


James L. Reveal! 


Professor Emeritus, University of Maryland 
College Park, Maryland 20742-5815, U.S.A. 
Honorary Curator, New York Botanical Garden 
Bronx, New York 10458-5026, U.S.A. 
jreveal@umd.edu 


ABSTRACT 


Six new combinations are required in Dodecatheon L. for a forthcoming treatment of the genus in Flora of North 
America north of Mexico: D. clevelandii Greene var. gracile (Greene) Reveal, D. clevelandii var. insulare (H 
Thomps.) Reveal, D. clevelandii var. patulum (Kuntze) Reveal, D. iene var. macrocarpum (A. Gray) Reveal 
D. pulchellum var. shoshonense (A. Nelson) Reveal, and D. utahense (N.H. Holmgren) Reveal. 


RESUMEN 


Se necesitan seis combinaciones nuevas en Dodecatheon L. en el tratamiento del género en la Flora of North 
America north of Mexico: D. clevelandii Greene var. gracile (Greene) Reveal, D. clevelandii var. insulare (HJ. 
Thomps.) Reveal, D. clevelandii var. patulum (Kuntze) Reveal, D. pulchellum vz (A. Gray) Reveal, 
D. pulchellum var. shoshonense (A. Nelson) Reveal, y D. utahense (N.H. ares Reveal. 


The following new combinations are required for a forthcoming treatment of Dodecatheon 
L.(Primulaceae) in volume 8 of Flora of North America north of Mexico. The rank of vari- 
ety is used throughout following Holmgren (1994). 
Dodecatheon clevelandii Greene var. gracile (Greene) Reveal, comb. nov. Dodecatheon See 
var. gracile Greene, Erythea 3:72. 1895. Type: U.S.A. CALIFORNIA. Santa Clara Co. gar 
specimen obtained from root collected at Loma Prieta by J.B. Davy (Not found). 


um 


f= 


Dodecatheon clevelandii Greene var. insulare (HJ. Thomps.) Reveal, comb. nov. Dodecatheon 
clevelandii subsp. insulare HJ. Thomps., Contr. Dudley Herb. 4:134. 1953 (as “insularis”). Type: U.S.A. CALI- 
FORNIA. Santa Barbara Co.: in chaparral along road to La Cumbre Lookout, 2.3 mi E of San Marcos Pass, 
Santa Inez Mts., 3000 ft., 25 Mar 1950, HJ. Thompson 1079 (HOLOTYPE: DS; ISOTYPE: OSC). 


pace clevelandii Greene var. patulum (Kuntze) Reveal, comb. nov. Meadia patula 
Kuntze, Revis. Gen. PL. 2:398. 1894. Dodecatheon patulum (Kuntze) Greene, Erythea 3:72. 1895. The earlier 
a name (Pittonia 1:211. 1888) is not valid as it was provisional when published. Type: U.S.A. CALIFOR- 
Co.: Livermore Valley, 23 Mar 1888, E.L. Greene s.n. (LECTOTYPE: NDG; ISOLECTOTYPE: DS), 

ae by HJ. Thompson, Contr. Dudley Herb. 4:135. 1953. 


The variation within Dodecatheon clevelandii, as defined by Thompson (1953), is main- 
tained with only the rank altered. The epithet var. gracile is equivalent to subsp. sanctarum 
(Greene) Abrams. 


Dodecatheon pulchellum (Raf.) Merr. var. macrocarpum (A. Gray) Reveal, oe nov. 
Do theon meadia L. var. macrocarpum A. Gray, Bot. Calif. 1:467. 1876. a U.S.A. ALASKA. Valdez- 
ordova Co.: Middleton Island, 2 Jun 1874, WH. Dall s.n. (LECTOTYPE: GH). N E: The ee pe was in- 
correctly termed a “neotype” by Beamish, Bull. Torrey Bot. Club 82:364. 1955. pee HJ. Thompson 

(1953) fad considered a “ BISve? specimen ( 27, GI Df rom the Santa Susana Mts., Ventura Co., Califor- 


considered t1 orm” of his var. macrocarpum, and 


nia, y 


annotated the prac eee “macrocarpum, igen atum.” 


This phase of Dodecatheon pulchellum has long been known as subsp. alas] (Hultén) 


‘Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, U.S.A. 


SIDA 22(2): 863 — 865. 2006 


864 BRIT.ORG/SIDA 22(2) 


Hultén or subsp. superbum (Pennell & Stair) Hultén. As treated by Thompson (1953) and 
by Taylor and MacBryde (1978), it is the tetraploid, coastal expression that occurs from 
southern Alaska to Oregon. 

Dodecatheon pulchellum (Raf.) Merr. var. shoshonense (A. Nelson) Reveal, comb. nov. 
Dodecatheon pauciflorum (Durand) Greene var. shoshonense A. Nelson, Bot. Gaz. 54:143. 1912 (as 
shoshonensis). TYPE: U.S.A. IDAHO. Twin Falls Co. Shoshone Falls, 26 Jul 1911, A. Nelson & J. F Macbride 
1362 (HOLOTYPE: RM). 

This variety of Dodecatheon pulchellum is found mainly in alkaline meadows on the foot- 
hills and valley bottoms of the northern Mojave Desert and Intermountain Region. In the 
latter area, var. pulchellum is a plant of the mountains. Only in the Pacific Northwest 
will one find var. pulchellum at lower elevations. The distinctive yellow pollen sacs of 
var. shoshonense are seen in populations of var. pulchellum in the Black Hills area of north- 
eastern Wyoming and adjacent South Dakota. Welsh (2003) proposed two entities which 
are synonyms of var. shoshonense: subvar. atwoodii and subvar. coriaceum. 


Dodecatheon utahense (N.H. Holmgren) Reveal, stat. nov. Dodecatl tatum Hook. var. utahense 
N.H. Holmgren, Brittonia 46:91. 1994. Type: U.S.A. UTAH. Salt Lake Co.: Moss Falls, Big Cottonwood Can- 
yon, Wasatch Range, 13 Jun 1977, N.H.H l g & PK. Holmgren 8412 (HOLOTYPE: NY, ISOTYPES: BRY, UTC). 

The work by Mastet al. (2001, 2004) demonstrated that varieties of Dodecatheon dentatum 

Hook., recognized by Holmgren (1994), should be regarded as distinct species. Accord- 

ingly three widely disjunct entities are now accepted: D. dentatum mainly on the eastern 

slope of the Cascade Range from south-central British Columbia to central Washington, 
with disjunct populations along the western portion of the Columbia River, the Colum- 
bia River Gorge of both Oregon and Washington, northeastern Oregon (Wallowa Co.), 
and in northern Idaho; D. ellisiae Standley of north central New Mexico in the moun- 
tains of Bernalillo, Sandoval and Torrance cos., and in southeastern Arizona in southern 

Apache, Greenlee, southern Graham, and northeastern Santa Rita cos.; and D. utahense,a 

narrow endemic confined to Big Cottonwood Canyon, Salt Lake Co., Utah. 

The recent molecular work on Primuldceae by Mast et al. (2001, 2004) and others 
(Kallersjé et al. 2000; Martins et al. 2003; Trift et al. 2002) has shown that recognition of 
Dodecatheon creates a paraphyletic Primula. Accordingly, Mast and Reveal (2006) trans- 
ferred all of the entities assigned to Dodecatheon to Primula. 


ACKNOWLEDGMENTS 


lam grateful to the Flora of North America Committee for providing funding to study 
Dodecatheon in the field and herbarium. To the several curators who made available their 
collections for study lam in your debt. In particular | must mention Kenton L. Chambers 
and Harry L. Thompson, both well acquainted with the genus, who willingly shared their 
decades of experiences. As always, the assistance of Kanchi Gandhi is indispensable. The 
Holmgrens, both Noel and Pat, provided me with information both from the literature 
and their experience with the genus. Both Kanchi and Noel formally reviewed the manu- 
script and lam grateful for their comments and timely corrections. Finally, my wife, C. 
Rose Broome, managed to survive the three months of getting Dodecatheon done; her 
help in herbaria and in the field is acknowledged with thanks. To the FNA editors and 
several reviewers who examined the “first draft” and provided comments and corrections, 
I give my thanks as we 


REVEAL, NEW COMBINATIONS IN DODECATHEON 865 
REFERENCES 


Beamish, K.. 1955. Studies in the genus Dodecatheon of northwestern America. Bull. Torrey Bot. Club 
82:357-366. 

HitcHcock, C.L. 1959. Dodecatheon L. Shooting star. Univ.Wash. Publ. Biol. 17(4):40-46. 

Hoimaren, N.H. 1994. Redefinition of Dodecatheon dentatum (Primulaceae) and rationale for use of 
varietial rank. Brittonia 46:87-94. 

KALLERSJO, M., G. Bercavist, and A.A. Anbersers. 2000. Generic realignment in primuloid families of the 
Ericales s.l: A phylogenetic analysis based on DNA sequences from three chloroplast genes and 
morphology. Amer. J. Bot.87:1325-1341 

Martins, L., C. OBeRPRIELER, aNd FH. Hetiwic. 2003. A phylogenetic analysis of Primulaceae s.l. based on 
internal transcribed spacer (ITS) DNA sequence data. Pl. Syst. Evol. 237:75-85. 

Mast, A.R., S. Ketso, A.J. RicHaros, D.J. LANG, D.M.S. Fetter, and E. Conti. 2001. Phylogenetic relationships in 
Primula L.and related genera (Primulaceae) based on noncoding chloroplast DNA. Int. J. PI. Sci. 
162:1381-1400. 

Mast, A.R., D.M.S. Fever, S. KeLso, and E. Conti. 2004. Buzz-pollinated Dodecath igi d from within 
the heterostylous Primula subgenus Auriculastrum (Primulaceae): A 7-region BcoENA phylogeny 
and its implications for floral evolution. Amer. J. Bot. 91:926-942. 

Mast, A.R. and J.L. Reveat. 2006. Transfer of Dodecatheon to Primula (Primulaceae). Brittonia, in press. 

THompson, H.J. 1953. The biosystematics of Dodecatheon. Contr. Dudley Herb. 4:73-154. 

Taytor, R.L. and B. MacBryoe. 1978. New taxa and nomenclatural changes with respect to Vascular 
plants of British Columbia:a descriptive resource inventory. Canad. J. Bot.56:184-195. 

Tritt, |, M. KALLERSJO, and A.A. AnperBerG. 2002. The monophyly of Primula (Primulaceae) evaluated by 
analysis of sequences from the chloroplast gene rbcL. Syst. Bot. 27:396-407. 

WELSH, S.L. 2003. Dodecatheon L. A Utah flora, ed. 3:526-527. 


866 BRIT.ORG/SIDA 22(2) 
Book REVIEW 

SUSANNA Ly_r. 2006. Fruit and Nuts. (ISBN 0-88192-759-7, hbk.). Timber Press Inc, 133 S.W. 

Second Avenue, Suite 450, Portland, OR 97204-3527, US.A.(Orders: wwwtimberpress.com, 


mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). $59.95, 
480 pp., color photos, figures, glossary, bibliography and index, 81/4" x 111/4". 


Have you ever looked at strange food in the grocery produce aisle and wondered which part you can eat? Or what 
preparation is needed to eat it? The new book Fruit and Nuts by Susanna Lyle will be a handy book to reference. 
Over two hundred food plants are presented in Fruit and Nuts along with details on floral structures, cultivation 
and uses. To help visualize the ae the book also includes over three hundred fantastic color photographs and 
ee 


Fruit and Nuts opens with a very informative section that introduces the readers to the various aspects of 
fruit and nuts and their production. Plant classification and re aan are covered in such a way that any 
reader can understand, and more detail can be found for bolded words in the glossary of the book. lyle gives 
readers botanical definitions of fruit and nut types with Sane examples and accented by color illustrations de- 
picting nut and fruit structures. Additional information in the introductory s 
by aa region, cultivation, propagation and the health benefits of nuts and fruits. 

e majority of the text is the fruit and nut “encyclopedia” in eer order by specific epithet from 
Acer sacc ae um (sugar my) to nics Pam as aebet should be noted that a few non-fruit nut items are 
included I t 


views of plant types 


1 fruit or nut entry contains a crisp color image of 
e edible portion of the plant along with descri oe structure, harvesting methods, cultivation, prop apation 
a other relative species. A side box for each 


ry lists uses such as food, nutritional value, alternative uses and 
ornamental value. The descriptions are very thorough and easy to read. An appendix covers plants for differing 
growth conditions/uses such as ornamental and health benefits. There is also a glossary, bibliography by topic 
and an index. 

Fruit and Nuts by Susanna Lyle is a fun and informative book with great value to those in food sciences, 
horticulture, gardening food plants or anyone with a general interest in learning more about the fruits and nuts 
consumed around the world. The text is thorough, easy to read, and is well illustrated with both color illustra- 
tions and color photographs. These illustrations and photographs are fantastic in helping readers to determine 

the edible portions of many Beale unknown fruits and nuts. This could be a wonderful reference book or 
gilt for cooks, horticultural students or “foodies” in general.—Lee Luckeydoo, Herbarium, Botanical Research In- 
stitute of Texas, 509 Pecan on Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(2): 866. 2006 


ERICAMERIA NAUSEOSA SUBSP. AMMOPHILA (ASTERACEAB), 
A NEW RABBITBRUSH FROM THE SAN LUIS VALLEY 
OF COLORADO 


Loran C. Anderson 
Professor Emeritus, ee ca ological Science 
tate Univer 
ihiestes FI fen pene U.S.A. 


ABSTRACT 


A previously undescribed, distinctive subspecies of rabbitbrush has been recently discovered in south-central 
Colotads ane it is ene dominant plant on eandenect = is e eequent on low sand aad It is = Bete natied 


ammophila. A 


r 


and a key is provided. 


RESUMEN 


Se ha descubierto recientemente una nueva subespecie de chamiso en el centro-sur de Colorado dinde es la 
planta commtanite en el arene) y eae en oe bajas. Se ha nombrado como Ered merida nauseosd subsp. 


ammophila p i y se aporta 


una Slave: 


One of the most widespread species in western North America is the rabbitbrush 
Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird. It ranges from southern Canada 
south to northern Mexico and from the Dakotas, Kansas, and Texas west to Washington 
and California. This rabbitbrush was long known as Chrysothamnus nduseosus (Pallas 
ex Pursh) Britton. H.M. Hall (with F E. Clements) monographed Chrysothamnus in 1923 
and Haplopappus in 1928, and his views held for many years. However, when the North 
American species of Haplopappus were demonstrated not to be closely related to the South 
American species (which included the type species for the genus), several sections of Hall's 
North American Haplopappus were recognized as separate genera. One such genus is 
Ericameria. Some species of Chrysothamnus, such as C. nauseosus, have long been noted 
for their close similarity to Ericameria. When DNA data suggested that the traditional 
Chrysothamnus was not monophyletic, Nesom and Baird (1993) transferred four of its 
species, including C. nauseosus, to Ericameria. 

Anderson (1986a) recognized 22 subspecies of C. nauseosus, whereas Nesom and Baird 
(1993) recognized two subspecies of E. nauseosa (ie., subsp. nauseosa and subsp. consimilis) 
and several varieties. Now, a new infraspecific taxon of E. nauseosa is to be described. 
Should it be a variety or a subspecies? Nesom and Baird (1993) emphasized the category 
of variety, but I believe my rationale for use of subspecies (Anderson 1986b) is solid, and 
I therefore describe the new subspecies, Ericameria nauseosa subsp. ammophila, below. 
Variability in the species is great, and my extensive study of the group in the field and in 
the herbarium has convinced me to apply infraspecific names to only the more signifi- 
cant morphotypes. It is surprising that this new subspecies has escaped recognition for 
so long. Specimens of it were not represented in the more than 12,000 collections of C. 
nauseosus that I studied from 56 herbaria (including COLO, CS, NCM, RM, and UNM) 
about 30 years ago. 


SIDA 22(2): 867 — 872. 2006 


868 BRIT.ORG/SIDA 22(2) 


intend to make appropriate subspecific combinations in Ericameria nauseosa ina 
future paper so that variation within the wider ranging subspecies can then be recog- 
nized through quadrinomials. Some argue that quadrinomials are cumbersome, but at 
an applied level, some field workers may wonder why the populations in their regions 
have the same subspecific name that is applied to somewhat different-looking popula- 
tions in other regions, and quadrinomials would help them see the relationships. There- 
fore, in the discussion section of the present paper, I will use subspecies names as they 
occurred in Chrysothamnus nauseosus (Anderson 1986a) for comparative purposes. 


ve nauseosa (Pallas ex Pursh) Nesom & Baird subsp. cual ue LG. een: 
nov. (Figs. 1, 2). Type: UNITED STATES. COLORADO. Saguache Co: dsheet 

ae a National Wildlife Refuge, ca. 4.5 air mi SW of Crestone, elev. 2528 m (7600 ft), Lat. 37° 56 12" N, Long. 
105° 44' 29" W, 2 Sep 2005, L.C. Anderson 21303 (HOLOTYPE: BRIT, IsoTyPEs: COLO, FSU, NY). 

Frutices usque ad 14 dm alti 2 I foliis {1 ibus; folia linearia, 20-45 

mi longa, 0.6-1.2 mm f] iculati itula cylindrica, 9.3-14 mm alta, straminea 

bracteis exterioribus ovatis, comentalosis vel glabratis, neous oblongis, glabris, apicibus obtusis vel 

acuminatis; ura geas 5, pallide flavi, corollibus 10.5-13 mm pone, lobis pecuaus unuequen lam longis, 

1] ] & I 


‘= 


aliquot y 

ion cterieis plerimane longioribus; achaenia aa. 
Fastigiate shrubs, 3.5-7(-14) dm tall, usually broader than tall from considerable branch- 
ing; ultimate flowering branches (8-)10-20(-25) cm tall, shallowly grooved, yellowish- 
green with compact tomentum, somewhat viscidulous. Leaves alternate, entire, linear, 
stiffly ascending or slightly spreading, (20-)25-30(-45) mm long, only slightly shorter 
toward stem tips, (0.6-)0.9-1.0(-1.2) mm wide, subterete with adaxial groove, light green, 
pubescence of sparse, compact, tomentum and a few longer, villous hairs. Inflorescences 
tightly congested paniculate cymes; heads cylindrical, (9.3-)11-12.5(-14) mm tall, (2-)2.5(- 
3)mm wide, stramineous; receptacle with central cusp (0.7-)1-2(-2.5) mm long. Phyllar- 
ies (15-)16-18(-20), in vertical rows, outermost (lowest) bracts ovate, 1.2-2 mm long, 
tomentulose to glabrate, inner bracts oblong, 8-10 mm long, glabrous with + ciliate mar- 
gins (few cilia to 0.4 mm long), weakly keeled, tips (rarely with tufted pappilae 0.1mm 
long) obtuse to acuminate. Disk flowers 5; corollas (10.5-)11.5-12.5(-13) mm long, nar- 
rowly cylindric gradually flaring to erect lobes (0.6-)0.7-LO(-L.3) mm long, yellow, sparse 
hairs on tube generally 0.2 mm long but a few 1-2(-3.5) mm long; styles (13-)14-16(-17.5) 
mm long, style branches 3.9-4.7 mm long, the stigmatic lines more or less equal to branch 
tips in length (40-56%), anthers 4 mm long with appendage 0.8-0.9 mm long; mature 
achenes cylindric, 5-nerved (the nerves when noticeable because of brownish material 
in associated secretory canals), +.5-6(-6.8) mm long, sericious with hairs (0.4-)0.5-0.7(- 
1.0) mm long, pappus (9-)10-11(-12.5) mm long, finely barbellate, white. 

Phenology.—A few shrubs of the new subspecies start to bloom in late August; most 
are in full bloom in mid-late September, whereas E. nauseosa subsp. consimilis, or var. 
oreophila (A. Nelson) G.L. Nesom & G.L. Baird, found in nearby more mesic, saline sites, 
blooms earlier in August. 

Habitat.—The new rabbitbrush occurs (in descending frequency) on: sandsheet, dune 
fields, stabilized sand dunes, and sandy soil of open pinyon-juniper (old dunes?). The term 
sandsheet is variously defined by geologists; here it refers to flat expanses of stabilized 
coarse, poorly sorted sands that occur extensively along the margins of the high dunes of 
the Great Sand Dunes National Park (tallest in North America). The rabbitbrush is the 
dominant plant (hiding nearly equally frequent grasses) on the sandsheet, and sandsheet 


LC / L oO I 1 


ANDERSON, ERICAMERIA NAUSEOSA SUBSP. AMMOPHILA, A NEW SUBSPECIES FROM COLORADO 


Fic. 1. Close-up of of flowering branches of Ericameria nausosa subsp.ammophila showing characteristic yellowish-green stems, nearly 
glabrous involucres, and short corolla lobes. 


870 BRIT.ORG/SIDA 22(2) 


rabbitbrush is very appropriate as a common name for this new subspecies (whose epi- 
thet ammophila means ‘sand loving’). From a distance, the vegetation on much of the 
Baca National Wildlife Refuge appears as a more or less evenly spaced, ‘unbroken sea’ of 
sandsheet rabbitbrush. Dune fields are generally flat but have some undulating topogra- 
phy. The sandsheet rabbitbrush occurs at elevations from 2518 to 2735 m (7550 to 8200 ft). 

Associated species.—The grasses, Achnatherum (Oryzopsis ) hymenoides (Roem. & 
Schult.) Barkworth and Chondrosum (Bouteloua) gracile H.B.K.,are usually co-dominants 
with the sandsheet rabbitbrush. The following are found at one or more sites: 
Chrysothamnus greenei (Gray) Greene, Eriogonum cernuum Nuttall, Heterostipa (Stipa) 
comata (Trin. & Rupr.) Barkworth, Ipomopsis longiflora Torr.) V. Grant, Nuttallia 
(Mentzelia) rusbyi (Wooton) Rydb., Opuntia polyacantha Haworth, Psoralidium 
lanceolatum (Pursh) Rydb., Rhus trilobata Nuttall, Senecio multilobatus Torr. & Gray, 
Sporobolus cryptandrus (Torr.) Gray,and Yucca glauca Nuttall. The flora of the sandsheet- 
sandhill ecosystem of this region is sparse in numbers of taxa. 


Additional collections: COLORADO. Alamosa Co.: Great Sand Dunes National Park Visitor's Center, 2 Sep 2005, 

. Anderson 21,280 (FSU); ie sheet by road to Medano Ranch, 9.9 mi E of Hwy 17, 2 Sep 2005, L.C. Anderson 
21,291 (FSU). Saguache Co.: ca. 3.7 mi SSE of Crestone, 21 Aug : 2001, J. Erdman O13 (FSU), 6 Sep 2001, J. Erdman 
O116 (FSU), 27 Sep 2001, J. Erdman 0121 (FSU); 1 Sep 2005, L.C. Anderson 21,276 (FSU); rd to San Isabel Cz ae 
trailhead, 21 Aug 2001, J. Erdman O1l4 (FSU); 7 Sep 2001 J. oud man O119 (FSU); 28 Sep 2001, J. Erdman 0123 (FS 
near Pinyon Flats, Great Sand Dunes National Park, 2 Sep 2 L.C. Anderson 21,286 (FSU); dune field near “Ox- 
bow Pond” between Spanish Creek and Cottonwood aie - 22 Aug 2001, J. Erdman O115 (FSU); 7 Sep 2005, J. 
Erdman O18 (FSU); 27 Sep 2001, J. Erdman 0122 (FSU), 2 Sep 2005, L.C. Anderson 21,301 (FSU). 


DISCUSSION 


The sandsheet rabbitbrush (subsp. ammophila) is perhaps most closely related to subsp. 
nitidus L.C. Anderson which occurs in northern Arizona and adjacent New Mexico. The 
two share the features of yellow-green foliage and stramineous, shiny involucres. They dif- 
fer in geography and habitat (subsp. nitidus prefers sandy gravels of dry stream beds) and 
in that subsp. ammophila fomrs shorter shrubs usually 35-70 cm tall (60-150 cm in subsp. 
nitidus), and has flowering stems fastigiate and more crowded than in subsp. nitidus, leaves 
usually 25-30 mm long and! mm or less wide (30-50 by 1-L5 mm in subsp. nitidus), corol- 
las mostly 11.5-12.5 mm long (9.5-11 mm in nitidus), unusually long trichomes on the 
corolla tubes and pubescent achenes (usually glabrous in nitidus). Subspecies turbinatus 
M_E. Jones is like the sandsheet rabbitbrush in height and branching, but it occurs gener- 
ally in sandy regions of the Great Basin and has mostly longer involucres with 
tomentulose phyllaries that are acute to acuminate (phyllaries in subsp. ammophila are 
mostly glabrous with obtuse to acute tips), and shorter corollas with lobes that are vil- 
lous. Subspecies consimilis (e., var. eae of Nesom & Baird 1993) rarely grows with 
subsp. ammophila, but subsp. consimilis usually grows in more saline or disturbed (mesic) 
habitats in this region, and it differs in forming generally taller, narrower shrubs with 
darker green foliage and smaller involucres and flowers. The unusually long hairs (up to 
m) on the corolla tubes of the sandsheet rabbitbrush are 10 times longer than any 
others found in the species or in all of the former genus Chrysothamnus (Anderson 1970). 
A portion of the Urbatsch et al. (2006) key to the varieties (listed here as subspecies) 
of Ericameria nauseosa is given below with the new subspecies included; subsp. 
ammophila shares a couplet with subsp. bernardinus, but the two are not closely related 
(ecologically, geographically, or taxonomically within the species). 


au 


AMARAQDUII A 


ANDERSON 


Fig.2.5 


Ad 
aun 


872 BRIT.ORG/SIDA 22(2) 


1. Cypselae a (plants 60-150 cm; involucres 10-12.5 mm) subsp. nitidus (in part) 
1. Cypselae hai 
16. Corolla ices villous (sometimes sparsely). 
19. Leaf blades 30-50 mm, faces glabrate subsp. nitidus (in part) 
19. Leaf blades 10-20 oe tomentulose subsp. turbinatus 


16. Corollas lobes glabro 
23. Phyllaries a epbeus outer sometimes sparsely hairy. 


24. Corollas 1 2[13] mm;involucres [9.3]10-14 mm. 
(24a. an lobes 1.7-2.3 mm subsp. bernardinus (in part) 
[24a. Corolla lobes 1 mm or less] subsp. ammophila 
24. Corollas 6—9+ mm; involucres 6-10 mm ___ subsp. consimilis [i.e., var. oreophila] 


ACKNOWLEDGMENTS 


James Erdman provided camaraderie, accommodations, transportation, and valuable 
general knowledge of the flora and geology of the area and obtained access for collecting 
in restricted areas. Richard Sparks of the USDA and Fred Bunch and Phyllis Pineda Bovin 
of Great Sand Dunes National Park also gave excellent assistance. Mark Garland provided 
the Latin diagnosis; William A. Weber and an anonymous reviewer offered important 
suggestions for improving the manuscript. Funds from the Friends of the Robert K. 
Godfrey Herbarium (FSU) helped defray publication costs of this paper. 


REFERENCES 


ANDERSON, L.C. 1970. Floral anatomy of Chrysothamnus (Astereae, Compositae). Sida 3:466-503. 

ANDERSON, L.C. 1986a. An overview of the genus Chrysothamnus (Astereae), In: E.D.McArthur and B.L. 
Welch. Proc. Symp. Biology of Artemisia and Chrysothamnus. USDA, Forest Service, Intermountain 
Research Station, Ogden, Utah. Pp. 29-45. 

ANDERSON, L.C. 1986b. Sympatric subspecies in Chrysothamnus nauseosus. In: E.D. McArthur and BLL. 
Welch. Proc. Symp. Biology of Artemisia and Chrysothamnus. USDA, Forest Service, Intermountain 
Research Station, Ogden, Utah. Pp. 98-103. 

Hatt, H.M. 1928. The genus Haplopappus; a phylogenetic study in the Compositae. Carnegie Inst. 
Wash. Publ. 389, Washington, D.C 

Hait, H.M.and F.E. Clements. 1923. The phylogenetic method in taxonomy. The North American spe- 
cies of Artemisia, Chrysothamnus, and Atriplex. Carnegie Inst. Wash. Publ. 326, Washington, D.C 

Nesom, G.L. and G.I. Baird. 1993. Completion of Ericameria (Asteraceae: Astereae), diminution of 
Chrysothamnus. Phytologia 75:74—93. 

Urearscu, L.E., L.C. ANDERSON, R.P. Roberts, and K.M.Neusic. 2006. The genus Ericameria Nuttall (Asteraceae). 
Fl. North Amer. 26:50-77. 


A NEW SPECIES OF BOLTONIA (ASTERACEAE) FROM THE 
RIDGE AND VALLEY PHYSIOGRAPHIC PROVINCE, U.S.A. 


John F. Townsend Vesna Karaman-Castro 
oe of Conservation and Recreatio Louisiana State University 
Division of Natural Heritage Department of Biological Sciences 
a7 ee St, 3° Floor 107 Life Science Building 
Richmond, Virginia 23219, U.S.A. Baton Rouge, Louisiana 70803, U.S.A. 


Email: John. Townsend@dcr.virginia.gov 


ABSTRACT 


Boltonia montana, a new f 1 i ine habi in the Appalachian Mountains of 
as New elie and Pennsylvania is described. This taxon has most often been es under the name 
onia roides (L.) LHer. due to its relatively large heads which are usually few in number, and vegetative 

] 


similarities. C 


of the new taxon, however, is dissimilar to that of B. oe and most closely 
resembles Boltonia ca relent na (Walt. ) Fern.,a species characteristic of coastal plain and Oia bottomlands 
of the Carolinas and Virginia. Boltonia montana differs from the latter taxon in terms of inflorescence structu 

head dimensions, ray color, phyllary morphology and size, plant stature, and leaf shape. Evidence of its eae 


tiveness is provided and its wetland habitat is discussed. 


RESUMEN 


omer | slA5 Ral . ¢ eee | “ J mak eS ci naae | 


ae i Vee Nee pas ui Pennsylvania ES taxon se ea eolecaas la mayor ce las veces con el 
usualmente en 


un numero pequeno, y hae vegetativas. La ene de las cipselas del nuevo Bon, sin embargo, es 
diferente de la de B. asteroides y se parecen mucho mas a Boltonia caroliniana (Walt.) Fern., una especie 
caracteristica de la llanura costera y base del pie de monte de las Carolinas y Virginia. Boltonia montana difiere 

e este Ultimo taxon en términos de estructura de la inflorescencia, dimensiones de la cabezuela, color de los 
radios, morfologia y tamano de los filarios, tamano de la planta, y forma de la hoja. Se aportan pruebas de su 
diferenciacion y se discute su habitat de zonas himedas 


INTRODUCTION 


The first author was alerted to the presence of a taxonomically problematic Boltonia in 
Augusta County, Virginia (Ridge and Valley physiographic province) by Gary P. Fleming, J. 
Christopher Ludwig (Virginia Department of Conservation and Recreation, Division of 
Natural Heritage), and Thomas F Wieboldt (Massey Herbarium, Virginia Polytechnic 
Institute) in late 2001. This plant had been identified as Boltonia asteroides by collectors, 
but certain details of the plants, particularly those of cypselae and phyllaries, indicated 
the Augusta County plants were not a good fit for that taxon. 

Subsequent investigations of the taxonomic literature, herbarium material, and living 
populations of Boltonia led to the realization that the plants from sinkhole ponds (dolines) 
in the Virginia mountains possessed a unique set of characters. During morphological 
and molecular anesrieaHens: of the nee plants, correspondence with - eos author 
led to the discovery of morphologically similar herbarium specimen dfrom the 
Ridge and Valley physiographic province of northwestern Nor iecee: and eastern Penn- 
sylvania. Field studies in the early 1980s and late 1990s by David Snyder and Kathleen 
Strakosch Walz of the New Jersey Natural Heritage Program uncovered or reinventoried 
the Boltonia populations in the northwestern part of that state and provided much data 


SIDA 22(2): 873 — 886. 2006 


874 BRIT.ORG/SIDA 22(2) 


on the plants discussed here. Subsequent herbarium and field investigations of the Vir- 
ginia, New Jersey, and Pennsylvania plants confirmed that these plants belonged to the 
same taxon. It is described here as new. 


Boltonia montana J.- Townsend & V. Karaman-Castro, sp. nov. (Figs. 1, 2). Typr: U.S.A. New 
Jersey. Sussex Co.: Muckshaw pond complex, located 3.2 km SW of Newton sual 14 km W of US 206, 
nearly dry, herb-dominated sinkhole pond, 3 Oct 2005, J.k Townsend 3570 (HOLOTYPE: US; IsoTYPES: MO, 
NY, VPD. 


Boltonia asteroides (L.) VHer. a oo maxime simile ob capitulis magnus et paucus sed acheniis glabris, 
7 ] R 


alis et pap] (Walt. Fern. similis. Phyllariis costa angustata, herbaceous 
et distalis viridis. 
Glabrous, perennial herbs 1.2-15 dm tall, ascending to erect (often prostrate in lower por- 
tion to ca. 1/2 stem length, sometimes rooting at nodes in this section). Stems glabrous, 
light to yellowish-green, ribbed with yellow or golden striations (more prominent in up- 
per 1/2 of plants), branching above into small, simple, corymbose panicles to diffusely 
paniculate in larger plants. Leaves basal and cauline, glabrous; basal leaves 0-18, alter- 
nate, in a rosette, oblanceolate to oblanceolate-spatulate, attenuated at the base (where 
often slightly widened), dark green or blue-green, often whitish or reddish colored at the 
base, with prominent, lighter-colored midrib and inconspicuous, arcuate secondary veins, 
apices bluntly rounded, mucronulate to entire, 1-17.5 cm long, 0.1-2 cm wide; cauline 
leaves oblanceolate to obovate or spatulate, bases sessile, often partially clasping, apices 
rounded to acute, mostly with mucronulate tips, appearing entire but minutely serrulate 
with narrow, wnlash, aoe margin, 2.7-11 cm long, 0.5-1.9 cm wide, sometimes 
with smaller leaves clustered at nodes. Capitulescences oes paniculate, few-headed 
(1-25) in average at many-headed (-100+) in large individuals. Peduncles bracte- 
ate, 0.5-11.8 cm long, 0.5-L1 mm thick, bracteal leaves oblanceolate to obovate or oblong, 
tips obtuse to acute, mucronulate, 0-5 per pedicel, 0.1-9.7 cm long, 0.02-1.4 cm wide. In- 
volucre 5.2-7.8 mm tall, 6.0-11.3 mm wide; phyllaries 30-00, lance-attenuate, serrulate to 
lacerate, in 2-4 series, 2.5-4.4 mm long, 0.4-0.9 mm wide, midrib 0.12-0.3 mm wide (1/10- 
1/5 of phyllary width), proximally whitish, distally green; receptacles conic, naked, L.9- 
3.0 mm tall, 1.9-3.8 mm wide. Ray flowers 27-406, pale lavender to pinkish (-white), corol- 
las glabrous, 4-5 nerved with golden or amber colored veins, 8.1-12 mm long, 0.8-1.9 mm 
wide, tubes 0.45-0.95 mm long; styles 1.3-2.45 mm long, style branches 0.25-0.4 mm long, 
cypselae glabrous, narrowly oblong, triquetrous, with | adaxial and 2 abaxial ribs, often 
slightly curved outward, 1.25-2.05 mm long, 0.5-0.9 mm wide, wings vestigial or absent, 
0-0.05 mm wide; pappus awns reduced to a crown of minute scales, 0-0.1(-0.4) mm long, 
Disk flowers 220-350, pale yellowish green to pale yellow-white as cypselae mature, su- 
ture lines pale amber, disk appearing more yellowish during anthesis, corollas glabrous, 
L.1-2.3 mm long, tubes 1.0-1.85 mm long, lobes 0.15-0.36 mm long; stamens included to 
exceeding corolla length by 0.4 mm, anthers 0.65-1.1 mm long, appendages 0.15 mm long, 
styles 1.05-2.55 mm long, style branches 0.2-0.4 mm long, cypselae glabrous, narrowly 
oblong, strongly flattened laterally with two marginal ribs, 1.3-1.9 mm long, 0.55-1.25 
mm wide, wings vestigial, O-0.09 mm wide, pappus awns usually reduced to a crown of 
minute scales 0-0.1 (-0.6) mm long. 

Distribution.—Distributed in sinkhole pond habitats and associated riversides and 
streamsides of the Ridge and Valley physiographic province in western Virginia, north- 
western New Jersey, and eastern Pennsylvania. 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A 


zk ‘ 
es Wh 


av 
I 


, 
} 


terial collected in $ Count y, New Jersey. 1 


C T. IOouUW. factrn Wl ra Pe L J 
hyll y ( t ).6 Ray corolla. 7. Disk corolla. Scale pareatale iif 


Fic. 
Habit. 2. Ray cypselae. 3. Disk cypselae. 4. Head. 5. Out 


except in 1 (= 1 dm) and 4 (=1 cm). 


876 BRIT.ORG/SIDA 22(2) 


Fic. 2. Low ing individual of Boltoni ith Eriocaul ticum, Augusta County, Virginia. Photo by Gary P. Fleming. 


I J 1 


Etymology.—The name refers to the species’ montane habitats. 

Phenology and development.—Flowering primarily occurs from early August to mid- 
October with precocious plants flowering in July, especially during dry years or in dr 
microhabitats. Hydrologic fluctuations in the species’ sinkhole pond habitat can ey a 
dramatic effect on the timing of flowering in a population. Even when water levels re- 
main high enough to inundate these ponds during the growing season, seeds are capabl 

of germination and rosettes may develop while still under water (Lassen and nee 
2003; David Snyder pers. comm. 2006) but the plants may not develop further, and if 
inundated for successive years the plants may die (David Snyder, pers comm. 2006). Au- 
tumnal plants produce basal offsets nearly identical in appearance to the rosettes pro- 
duced by young plants. In the case of reclining or ascending plants of Boltonia montana, 
adventitious roots may form at the nodes as autumn approaches. In many cases, these 
nodes also produce leafy offsets complete with root systems. During collection of flower- 
ing or fruiting plants, these vegetative propagules are often dislodged due to their weak 
attachment to the parent plant. 

A concentric gradient of environmental conditions is created in these sinkholes by 
fluctuating hydrology and sloping shorelines. It is therefore possible to find several co- 
occurring growth stages of Boltonia montana. Recent field work in New Jersey with Natu- 
ral Heritage Program Botanist David Snyder and Ecologist Kathleen Strakosch Walz (au- 
tumn 2005) was conducted following a long period of pond inundation. On the pond 
margins that were not recently inundated, relatively well-developed plants (with flow- 
ers, fruits, and basal offsets) were found. In more recently inundated portions of the ponds, 
large numbers of Boltonia rosettes were found. These plants were either newly germi- 


7 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 877 


nated seedlings, rosettes persisting from a previous germination event, or vegetative off- 
sets now independent of the parent plant. Regardless of origin, these rosettes were unable 
to flower or proeuce fruit before the end of the growing season due to their ey ceccpuonally 
late Septem ber —October). In general, when f as ground- 
water levels drop after such periods of inundation, new ae develop fro rom the seed bank 
(David Snyder, pers. comm. 2006). 

Observations indicate that Smith and Mettler’s (2002) description of the life cycle of 
the Federally Endangered Boltonia decurrens (Torr. & A. Gray) Wood provides a frame- 
work for understanding the life cycle of Boltonia montana as well: “Although B. decurrens 
is a perennial under optimal conditions, various life stages may behave as winter annu- 
als (winter seedlings bolt, flower, and senesce the following summer), as summer annu- 
als (summer seedlings bolt, flower, and senesce the same summer), or as biennials (ca. 25 
percent of spring seedlings overwinter as rosettes that bolt, flower, and senesce the fol- 
lowing summer).” Such a patchwork of potential strategies makes sense for a species 
adapted to the highly variable flood pulse of Midwestern rivers (B. decurrens) as well as 
the dramatic hydrological fluctuations of isolated wetland systems (B. montana). 


HABITAT AND DISTRIBUTION 


Boltonia montana is known from localized sections of the Ridge and Valley physiographic 
province in three U.S. states: Virginia (one county), New Jersey (two counties), and Penn- 
sylvania (one county). All occurrences are found within areas of gently sloping, rolling, 
or nearly level ground between 149 and 488 m (490-1600 ft) altitude (Walz et al. 2001, 
Va. Division of Natural Heritage files), but many of the New Jersey ponds are surrounded 
by rough terrain with extensive bedrock outcrops (Kathleen S. Walz, pers. comm. 2006). 

The sinkhole pond complexes which support this species span an area measuring 
roughly 14x 15 km near Newton, New Jersey and 5.25 x 4.5 km near Waynesboro, Vir- 
ginia. Within these areas, plants are known from a total of 36 sinkhole ponds and lakes 
(22 in New Jersey and 14 in Virginia; Va. Division of Natural Heritage files; David Snyder, 
pers comm. 2006). Although the vast majority of the records for this new taxon come 
from isolated wetlands, a very few have been found along rivers. Besides two collections 
made the same day in 1936 along the South River in Virginia, the only other such collec- 
tions were made in 1864 (1 specimen) and 1865 (1 specimen) along the Susquehanna River 
in Dauphin County, Pennsylvania—apparently the only observations of this species in 
Pennsylvania. The Virginia occurrence was taken from an area extremely close to, and 
just downslope from, a sinkhole pond complex known to contain Boltonia montana, while 
the disposition of the Pennsylvania material relative to potential sinkhole pond seed 
sources is unclear. It is assumed here that only occasional plants become established in 
such riverine habitats from nearby “parent” populations. The lack of additional collec- 
tions at least indicates that such riverine habitats are rarely occupied. 

Like Boltonia montana, many species seemingly defined by their isolated wetland 
habitat are occasionally found in other community types, indicating that their affinity 
for these ponds is not restrictive. Examples include the co-occurring Helenium virginicum 
Blake in Virginia, which is almost entirely restricted to sinkhole ponds but also occurs in 
a wet meadow of the nearby South River (Va. Division of Natural Heritage files). The 
diminutive sedge Fimbristylis ee eae ex Small and Britt.was also thought to 
bea strict pond endemic and func hin most localities, but discoveries in North 
and South Carolina (Leonard 1981. 1987), established the species as an occupant of river- 


878 BRIT.ORG/SIDA 22(2) 


ine “draw-down” bars, banks, and sloughs as well. A final example involves Boltonia 
decurrens, a perennial species of floodplains in the Ilinois River valley (Stoecker et al. 
1995) which was originally dependent on natural flood pulses for habitat maintenance 
and germination. Most of the habitats occupied by this species have been described as 
wet prairies or bottomland communities of various types, but McClain et al. (1997) also 
reported the species from “sand ponds” near the Illinois River in Cass and Mason coun- 
ties, Illinois. They described water levels in the ponds as fluctuating in response to the 
local aquifer rather than overbank flooding, but it is not known to what degree human 
manipulation of the river has lessened the importance of this water source in the ponds. 

The sinkhole pond habitats occupied by Boltonia montana are ultimately formed by 
the solution of underlying carbonate bedrock and subsequent volume loss or collapse of 
this foundation (Fleming @ Van Alstine 1999; Walz et al. 2001). The habitat types found 
within sinkhole ponds in Virginia and New Jersey are floristically and physically differ- 
ent, owing to their means of origin, Pleistocene history (glacial scouring and surficial 
geologic deposits), and distance of calcareous materials from the soil surface. 

In Virginia, weathering of limestones and dolomites of the Cambrian Shady and 
Elbrook Formations is ultimately responsible for the formation of these pond features, 
but a deep overburden of relatively acidic, infertile soils covers these carbonate rocks toa 
depth of 30 to 150m (Fleming & Van Alstine 1999). This phenomenon is similar to that 
described in Maryland and West Virginia by Bartgis (1992) who studied ponds support- 
ing the rare pond endemic Scirpus ancistrochaetus Schuyler. In Virginia, surficial depos- 
its are the result of alluvial fan formation along the western slopes of the Blue Ridge 
mountains. Materials range from quartzite cobbles and boulders to sand, silt, and clay 
and range from the late Pleistocene to Pliocene or Miocene in age (Whittacar et al. 1999). 
Due to the depth where the carbonate rocks reside, soil and water chemistry of the ponds 
is acidic (pH 3.9-4.0 at 0-30 cm). Soils are predominantly gray to blackish clays or clay 
loams with lighter colored mottling evident below 3 to 10 cm. This clay, weathered from 
metasiltstone and phyllite alluvium, impedes drainage but most ponds draw down as 
the growing season progresses and evapotranspiration increases. Although some of these 
sinkholes are relatively stable hydrologically, Boltonia montana is almost exclusively 
found in those ponds with highly variable and unpredictable water levels. The Boltonia 
montana “zone” in these ponds is typically doughnut-shaped and extends from the rim 
of the pond to the edge of the most deeply flooded portion of the basin (Fig. 3). 

The y types occupied by the species in Virginia are the Quercus palustris / 
Panicum rigidulum - Panicum verrucosum ~ Eleocharis acicularis Wooded Herbaceous 
Vegetation Association and the Cephalanthus occidentalis / Polygonum hyd ropiperoides 
- Glyceria acutiflora - Proserpinaca palustris Shrubland Association (Grossman et al. 
1998; NatureServe 2005). The former community type is endemic to Virginia and assigned 
a rarity rank of G1G2 (NatureServe 2005). The latter type is somewhat more widespread 
(ranked G3?/S1) but is considered endemic to mid-Atlantic region. These communities 
are open-canopied except on pond edges, where the forest canopy can provide partial 
shading. Associated species in these communities include Quercus palustris, Nyssa 
sylvatica, and Cephalanthus occidentalis (mostly occurring on pond margins), Panicum 
rigidulum var. rigidulum, P. philadelphicum, P. verrucosum, Eleocharis acicularis, 
Eleocharis melanocarpa, Eriocaulon aquaticum, Lysimachia hybrida, Ludwigia palustris, 
Leersiaoryzoides, Hypericum boreale,and the federally Threatened Helenium virginicum. 
In addition to relatively undisturbed sinkhole ponds, Boltonia montana is also found with 


— 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 879 


Fic. 3. Sinkhol 1 habi f Roltoni, i Augu ta County. Virdinia. Photo by G oc 
J y 


dry Fr. PIecninniy. 


Fig. 4.2 ied by Boltoni | in of “Piggyback Pond,” Warren County, New Jersey. Photo by John F. Townsend. 


Pr 7 J J 


880 BRIT.ORG/SIDA 22(2) 


H. virginicum in several altered ponds, most of which are in pasture. As a result, these 
ponds include a number of species with broader habitat associations such as Juncus effusus, 
J. canadensis, J. tenuis, Bidens frondosa, B. polylepis, Ambrosia artemesiifolia, Phytolacca 
americana, Daucus carota, Vernonia noveboracensis, Digitaria ischaemum, Echinochloa 
crusgalli, Datura stramonium, Eclipta alba,and Erigeron annuus,among others. 

The New Jersey ponds are decidedly calcareous, occurring within a glaciated karst 
landscape on dolomite bedrock. These sinkholes are found within the Ordivician Allen- 
town Dolomite and Beekmantown Dolomite formations, with surficial deposits composed 
of a glacial till/ bedrock complex. In rare instances, Boltonia montana populations may 
be found on glacial outwash (kettles). In all cases, the amount of overburden present is 
significantly less than (and more alkaline than) that found in the Virginia pond com- 
plexes, and the carbonate rocks or marl deposits are usually at or near the soil surface, 
providing a slightly acidic to mildly alkaline soil environment (pH 6.2-7.8 at 0-30 cm; 
Walz et al. 2001). In New Jersey the presence of marl and Chara spp. are characteristic of 
the sinkhole pondshore habitat for B. montana. Although soil chemistry of these ponds 
differs from Virginia examples, dramatic fluctuations in the seasonal water table (as much 
as 5.5m per season in New Jersey) are characteristic of ponds occupied by the species in 
both states. Soils in New Jersey are shallow silt loams, silty clay loams, silty clay, or rarely 
silt marl overlaying clay or bedrock. Marly muck or peat soils are also present in rare in- 
stances and small dolomite rocks are often found on the surface (Walz et al. 2001). Deep 
marl deposits often occur in the center of ponds occupied by B. montana, but as with the 
Virginia populations, Boltonia montana usually occurs ina distinctive doughnut-shaped 
zone or “ring” between the pond edge and the more frequently and deeply flooded pond 
center (Fig. 4). 

The community type occupied by the new taxon in New Jersey is the Boltonia 
asteroides var. asteroides - Symphyotrichum racemosum - Mentha arvensis Herbaceous 
Vegetation Asociation (Grossman et al. 1998; Natureserve 2005; Walz et al., 2001), a com- 
munity type that is ranked G1G2 by Natureserve (the naming of B. montana will involve 
a concomitant change in the community name). Characteristic species in this commu- 
nity type include: Asclepias incarnata, Cyperus strigosus, Carex cryptolepis, C. lurida, C. 
typhina, C. viridula, Cuphea viscosissima, Eleocharis palustris, Eragrostis frankii, Eupa- 
torium perfoliatum, Iris versicolor, Dichantheliumacuminatum var. fasciculatum (=Pani- 
cum lanuginosum var. tennesseense), Panicum capillare, Panicum rigidulum var. pubescens 
(=Panicum longifolium), Polygonum amphibium, Sium suave, the green algae Chara spp., 
and the bryophytes Hypnum lindbergii, Campylium stellatum,and A serpens 
(Walz et al. 2001). Species in this community type which are rare in New Jersey include: 
Boltonia montana, Carex bebbii, C. cryptolepis, C. haydenii C. retrorsa, C. typhina, C. 
viridula, Eleocharis quadrangulata, and Eragrostis frankii (Snyder 2005). 

Many endemic wetland plant species are confined to habitats with widely fluctuat- 
ing hydroperiods, and it has been suggested that the ability of these species to tolerate 
such stressful conditions releases them from competition with other taxa (Knox 1997). 
This aspect of the life history of Helenium virginicum was studied by Knox (1997), who 
concluded that the ability of plants to tolerate the twin stressors of potentially toxic soils 
and fluctuating hydrology was rare and contributed to the species’ endemism. Boltonia 
montana and H. virginicum co-occur at several Virginia ponds where these adverse soil 
conditions (low pH and elevated levels of aluminum and arsenic) have been documented, 
with the remainder of Virginia ponds thought to have similar soil characteristics. Data 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 881 


on soil relations in the New Jersey ponds are incomplete, but indications of high alumi- 
num, calcium, and magnesium levels have been found and erratic water level fluctua- 
tions are well-documented there (Walz et al. 2001). 


Additional specimens examined: NEW JERSEY: Sussex Co.: near Lake Grinnell, 17 Sep 1887 N.L. Britton, s:n(NY); 
Swartswood Lake, 6 Jul 1907, Philip Dowell 4951 (GH), low swale, Brighton, 4 Sep 1910, K.K. Mackenzie 4778 (GH, 
NY); borders of pond E of Swartswood Lake, Aug 1911, K.K. Mackenzie 4922 (NY, PH); border of Muckshaw Pond, 
Springdale, Aug 1917, K.K. Mackenzie 7986 (NY); Ea ae Pond,” 3.1 km E of Swartswood, 4.7 km NW of 
Newton, 3 Oct 2005, J.F Townsend 3556 (VPI, NCU, CLEMS); “Duck Pond,” Swartswood State Park, 1.2 km N of 
Paulinskill, 5 km WNW of Newton., 3 Oct 2005, J.F ay nsend 3557 (PH, PENN); “Frog Pond,” Swartswood State 
Park, 1.75 km NNE of Paulinskill, 5 km WNW of Newton., 3 Oct 2005, J.E Townsend 3558 (NY); “Little Frog Pond,” 
Swartswood State Park, 1.75 km NNE of Paulinskill, 5 km WNW of Newton, 3 Oct 2005, J.F Townsend 3559 (VPI); 
“Spring Lake,” Swartswood State Park, 3.1 km NW of Newton, 2.1 km SE of Swartswood, 4 Oct 2005, J.E Townsend 
3562 (BRIT, GMUF GH); northernmost “Muckshaw Ponds,” 1.6 km SW of Newton and 0.4 km W of US 206, 4 Oct 
2005, J.E Townsend 3565 (MARY, WYVA, VPI): “Muckshaw Ponds.” 2 km SW e Newton and 0.6 km W of US 206, 4 
Oct 2005, J.F Townsend 3566 (CHRB, PH, NY); “Muckshaw Ponds,” 2.75 km SW of Newton and 0.6 km W of US 
206, . Townsend coe NCU, GMUF). Warren Co.: Muddy calcareous ae and flats, Shyster Pond, 24 Jul 
1920, K.K. Mackenzie Y, PH); muddy shore of limestone sink pond, eee 4 Oct 1981, DB. Snyder 
551-6P - ee Pig ac ee located ca. 5 km NE of Blairstown and 1.6 km N of Squires Corner, J.E Townsend 
3560 (GH, CLEMS, USCH); (Warren/Sussex Co. line) “Four-Angle Pond,” fae 16 km NW of Tranquility and 
1.25 km a W om es, 3 Oct 2005, J.F Townsend 3561 (VPI). PENNSYLVANIA. Dauphin Co.: banks of the 
Susquehanna, near Dauphin, 2 Aug 1865, CE Parker, s.n. (NY): Harrisburg, Jul 1864, LC. Martindale, s.n.(PH). VIR- 
GINIA. Augusta Co.: Sandy, wet soil, bank of South River, vic. Lipscomb, 18 Jul 1936, L.L. Carr 47 (GH); growing on 
bank of South River vicinity of Lipscomb, abundant in Stuart's Draft area around ponds and along South River, 
18 Jul 1936, E.S. Rawlinson 58A (VPP: Kennedy Mt. Meadow, (unintelligible) bog, 14 Sep 1937, L.L. Carr, s.n. (PH); 
moist edge of pond (open sit.), vic. Lipscomb, 5 Aug 1938, L.L. Carr 365 (GH); edge of dried up pond, Big Levels 
refuge, 8 Sep 1939, A.B. Massey, s.n. (VPI); Lipscomb Pond, in water, vicinity of Lipscomb, Shenandoah Valley, 8 Sep 
1940, EP Killip, 36069 (US); adjacent to Spring Pond, near Stuarts Draft, bottom of dried up pond, 16 Sep 1964, R. 
S. Freer Sete (GH, ve ieee to Spring Pond, near Stuarts Draft, bottom of dried up pond, 23 Sep 1964, R.S. 
Freer 2869 (VPD); edge of pond near Spring Pond, 5 Aug 1967, A.M. Harvill 17240 (FARM, NCU); Twin Ponds, 
near Sherando, common in aca portions of pond, elev. 1550 ft, 19 Sep 1970, CE. Stevens 2781 (VPI); dried up 
pond, open area, Twin Ponds, Big Levels Game Management Area, 19 Sep 1970, LE Wieboldt 271 (WILLD; around 
the margins and drawdown areas of Spring Pond, elev. 1600 ft, 26 Sep 1970, G. Ramsey, R.S. Freer, R. Bruce, et al. 
18260 (WILLD; common in shallow water of open edge of Twin P Pond, near Sherando, elev. 1540 
ft, 16 Oct 1971, CE. Stevens 4772 (VPI, WILLD; Wood duck Pond, scattered in aod around edges of shallow es 
hole pond 1.2 mi SE of Lipscomb, elev. 1440 ft, 2 Oct 1984, C.E. Stevens 19078 (FARM); Campbell’s Pond, 1.7 mi N 
of Sherando, SW side of St. Rt. 634 1 mile N of jet. with St. Rt. 610, elev. 1470 ft, 6 Aug 1986, G. Fenwick, s.n. aD 
common with Eriocaulon septangulare in dried muddy bed of intermittent sinkhole pond (“Twin Pond”), 2.3 mi 
W of Sherando, elev. 1540 ft, 22 Sep 1991, G.P. Fleming 6126 w/ CE. Stevens (WILLD; Twin Ponds, located ca. 2.26 
mi due W of Sherando, 8 Aug 2002, J.K Townsend 2869 (VPI, WILLI, Va. Div. Natural Heritage Herbarium). 


Relationship to Boltonia asteroides var. asteroides and B. caroliniana 

The gross morphology of Boltonia montana is nearly identical to that of B.asteroides var. 
asteroides, a fact which explains the use of this name in both Virginia and New Jersey for 
many years. Boltonia asteroides var. asteroides will be referred to as B. asteroides for the 
sake of brevity in the following discussion. 

The cypselae of Boltonia montana are found to lack several key characteristics found 
in B. asteroides, namely: cypselae wings, cypselae trichomes, and pappus awns. Occasional 
plants of B. montana will posess cypselae with slender pappus awns that are longer than 
the typically low crown of scales. In this respect, such plants may have awns approach- 
ing the length seen in B. asteroides, but the other key features, glabrous cypselae with 
wings absent or vestigial, readily separate them from B. asteroides. Fernald (1940) de- 
scribed B.asteroides as having obovate, thick rimmed cypselae with awns wanting or up 
to 0.7 mm long and apparently included the plants of northwestern New Jersey in his 


882 BRIT.ORG/SIDA 22(2) 


concept of B.asteroides at that time, which probably accounts for his description of awn- 
less B. asteroides in Gray’s Manual (Fernald 1940, 1950). In many specimens of B. asteroides, 
phyllaries are mostly sclerified and whitish on either side of the midrib and almost to 
the phyllary tip, becoming greenish gradually in the distal portion, and have a relatively 
large, resinous midrib (1/5 to 1/4 of phyllary width) which continues to the phyllary tip. 
In contrast B. montana phyllaries are only sclerified and whitened in their lower half or 
less, becoming green and herbaceous in the distal portion. The midrib is relatively nar- 
row (1/10 to1/5of phyllary width) in the lower half and anastamoses in the distal half to 
produce a network of fine veins within the green portion of the phyllary. It was at first 
assumed that the phyllaries seen in B. montana were always unlike those found in B. 
asteroides, but several specimens of B. asteroides from the Susquehanna River in Pennsyl- 
vania have similar phyllary morphology to the former. Whether this represents past gene 
exchange with B. montana or character convergence is not known. 

Due to the strong resemblance between the cypselae of Boltonia montana and B. 
caroliniana, the latter species was investigated to discern other morphological similari- 
ties, if present. Other than cypselae morphology, these two taxa have few character states 
in common. 

Phyllaries of Boltonia caroliniana are quite dissimilar to B. montana, and most closely 
resemble B. asteroides in terms of their white color and relative midrib size, but are shorter 
and narrower than both B. asteroides and B. montana. Head size, ray corolla length, and 
pedicel diameter of B.caroliniana are smaller than in B. asteroidesand B. montana, giving 
a delicate appearance to the inflorescence of B. caroliniana. In addition, typical individu- 
als of B. caroliniana are profusely branched and have numerous heads. Most individuals 
of B. montana and B. asteroides have relatively few heads (often 1-25), although plants of 
both species have been noted to produce large plants with numerous heads (Karaman- 
Castro and Urbatsch 2006, see below). Such large reproductive outputs are seldom seen 
and may be a strategy to sidestep adverse meterological or edaphic conditions. Ray corol- 
las of B. caroliniana, in addition to being smaller than those of B. montana, are white, 
fading to pink or lilac in age (Anderson 1987, pers. obs.), while the rays of B. montana are 
usually lilac to pinkish throughout the flowering period, though they may also be extremely 
pale (see Fig. 2). Most plants of B. montana are characterized by widely oblanceolate leaves 
whereas those of B. caroliniana are linear to narrowly oblanceolate or linear-obovate. This 
difference is most evident when bracteal leaves of the two species are compared. 

Morphological similarity of Boltonia montana to B. asteroides and B.caroliniana sug- 
gest that B. montana may have arisen as a result of an ancient hybridization between the 
two species. Phylogenetic analyses of the internal and external transcribed spacers (ITS 
and ETS) from the nuclear ribosomal DNA suggested sister relationaship of B. montana 
and B. caroliniana, while only distant relationship with B. asteroides (Karaman-Castro et 
al. in prep.). Thus the treatment of the awnless and wingless specimens of B. asteroidesas 
a new species, B. montana was supported by molecular data. However, it remains unclear 
whether B. montana isa result of an ancient hybridization between B. caroliniana and B. 
asteroides. 


Some observations on the morphology of Boltonia montana 

The stature of Boltonia montana varies from ascending to erect, with most individuals 
exhibiting the former condition (Fig. 1). It seems probable that this growth pattern is a 
response to edaphic conditions in the ponds rather than genetics. For instance, observa- 
tions at “Twin Ponds” in Augusta Co., Virginia were generally of few-flowered, ascending 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 883 


or erect plants from 2 to 6 dm in vertical extent (Fig. 2). This growth form is present in 
most populations, but was completely absent during the 2002 season at this site, where 
all plants were erect, .5-2.0 m tall, bore numerous heads, and had narrower bracteal leaves. 
Since this growth form was produced at the height of a multi-year drought, it seems pos- 
sible that soil moisture relations were involved in mediating this morphological shift. 
Herbarium specimens and observations of New Jersey populations also produced the 
general impression of decumbent or low growth, but plants in somewhat drier micro- 
habitats tended to be taller, more strictly erect, have more heads, and possess narrower 
bracteal leaves. In light of this variability, the particulars of inflorescence size and shape, 
head distribution, and plant stature should not be considered diagnostic of this taxon. 
Adventitious roots sometimes form at nodes along the decumbent portion of stems 
in Boltonia montana. These roots are either associated strictly with the stem or may be 
attached to leaves similar in appearance to basal rosettes. These “offsets” are a means of 
vegetative propagation and are often easily detached from the plant in the fall of the year. 
Such rooting along the stem was encountered more often in New Jersey populations than 
in Virginia localities and basal offsets were produced in all populations studied. These 
structures are hypothesized to be the primary means of perennation in the species. 
Smith and Keevin (1998) describe the adaptations for long-distance water dispersal 
in Boltonia decurrens cypselae and pointed to the evidence of such dispersal under field 
conditions. They found the wide, flat profile of the cypselae and wings, along with the 
trapping of air bubbles by cypselae trichomes, allows them to easily rest on the surface 
of the water. In one experiment, 20% of cypselae were still floating after + weeks of simu- 
lated wave action. No similar experiments have been done with B. montana, but cypselae 
morphology implies a reduced level of dispersability when compared to all other mem- 
bers of the genus except B. caroliniana. Given that nearly all B. montana populations are 
found in small, island-like wetlands within an upland matrix, these physical character- 
istics may have developed due to the improved survival of sedentary (wingless, awnless, 
hairless) cypselae. Schiffman (1997) noted wing width reductions in cypselae from is- 
and populations of Coreopsis gigantea (Kellogg) Hall when compared to conspecific 
mainland individuals. She theorized that this atrophy occurs because “dispersal-enhanc- 
ing morphologies increase probabilities of being transported beyond the bounds of these 
narrow habitats,” a classic argument applied to many specialized taxa of islands. In the 
case of Boltonia montana, the reasons for its restriction to such narrow “island-like” habi- 
tats are only speculative but it seems likely that a competitive advantage is enjoyed by 
the species in sinkhole ponds and enhanced dispersal structures would represent an un- 
necessary expenditure of energy. Occasional overland flows do provide occasional con- 
nections between neighboring sinkhole ponds in Virginia, (Christopher S. Hobson 2005, 
pers. comm.) and despite lacking obvious dispersal mechanisms, B. montana propagules 
could conceivably disperse to adjacent ponds during these relatively rare overflow events. 


A KEY TO BOLTONIA MONTANA AND RELATED SPECIES 


Given the morphological variability observed in Boltonia species, several morphological 
characters should always be used when identifying these taxa. Taken together, the fea- 
tures of phyllaries, and cypselae awns, wings, and surfaces are diagnostic of Boltonia 
montana. 

The following key distinguishes Boltonia montana from B. asteroides var. asteroides 
and B. caroliniana, the two congeners most likely to be confused with the new taxon. 


884 BRIT.ORG/SIDA 22(2) 


1. Cypselae with two distinct pappus awns in addition to a shorter ring of scales, the awn stly 
0.8-1.1mm long; cypselae wings obvious, mostly 0.3-0.4mm wide; lateral faces of cypselae 
pubes Boltonia ast Jes var. asteroides 


ent 
; seer with pappus reduced to a short ring of scales to 0.15mm long or with occasional 
slender awns to 0.6mm; cypselae wings lacking or up to 0.1mm wide; lateral faces of cypselae 
glabro 


—_ 


N 


ee 0.2-0.5mm wide, sclerified throughout or with herbaceous a midrib relatively 

prominent (1/5-1/3 phyllary width); heads 4.3-6.1mm wide, peduncles 0.25-0.5 mm in di- 

ameter;ray corollas white fading to pinkish in age; cauline and bracteal a narrowly oblan- 
ceolate to linear Boltonia caroliniana 

Phyllaries 0.4—0.9mm wide, sclerified in lower 1/3 to 1/2 distal portion dilaae mid- 

rib relatively narrow (1/10-1/5 phyllary width);heads 6-11.5mm wide, peduncles 0.5—1.1 mm 
in diameter; ray corollas lilac or pinkish (-white); upper leaves oblanceolate to oblong Boltonia 
montana 


N 


CONSERVATION 


Isolated wetlands of the eastern United States are known for harboring a large number of 
rare species and for being heavily impacted by human activity (Weakley & Schafale 1994; 
Edwards & Weakley 2001; Comer et al. 2005). Research has focused heavily on depres- 
sion wetlands of the coastal plain, but some studies have dealt with rare and disjunct 
species affiliated with depression wetlands of the interior (Steyermark 1963; Homoya 1983: 
Bartgis 1992; Fleming & Van Alstine 1999), including the rare pond endemics Helenium 
virginicum and Scirpus ancistrochaetus. 

Ponds containing Boltonia montana have been the focus of conservation efforts in 
both Virginia and New Jersey due to the large number of globally and regionally rare 
plants and animals already known to occur in them. A patchwork of private and public 
ownership exists for these ponds, and the level of threat to these wetlands is generally 
high due to their small size and location within upland areas. A recent Supreme Court 
ruling also called into question the jurisdiction of the Clean Water Act over isolated wet- 
lands (Comer et al. 2005), potentially relaxing protections for these natural ponds and 
their characteristic flora. Given their vulnerability and relative scarcity on the landscape, 
efforts to protect these unique wetlands should continue to be a high priority. 


ACKNOWLEDGMENTS 


We thank staff of the Virginia Department of Conservation and Recreation, Division of 
Natural Heritage for their insight and support during this project. Thomas F. Wieboldt of 
Virginia Polytechnic and State University provided taxonomic expertise and access to 
specimens. David B. Snyder of the New Jersey Natural Heritage Program and Kathleen 
Strakosch Walz of the New Jersey office of the Nature Conservancy provided detailed 
collections data, maps, field expertise, and interpretation of the ecological communities 
and karst landscape of northwestern New Jersey. Greg Plunkett of Virginia Common- 
wealth University coordinated loans from various herbaria and provided work space. The 
staff of the Gray Herbarium, Harvard University (GH), the University of North Carolina- 
Chapel Hill (NCU), Longwood University (FARM), and the College of William and Mary 
(WILLD are thanked for their hospitality while studying specimens. Karen Townsend is 
thanked for scanning and manipulating the illustrations comprising Figure 1. Alan 
Weakley and an anonymous reviewer provided valuable suggestions for improving the 
manuscript. Special thanks go to Adam and Zena Rudzki for access to, and permission to 
collect, the type voucher specimen from their private property in New Jersey. 


TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 885 


REFERENCES 


ANDERSON, L.C. 1987. Boltonia apalachicolensis (Asteraceae): A new species from Florida. Syst. Bot. 12: 
33-138 

Bartais, R.L. 1992. The endangered sedge Scirpus ancistrochaetus and the flora of sinkhole ponds in 
Maryland and West Virginia. Castanea 57:46-51. 

Comer, P., K. Goopin, G. Hammerson, S. MENARD, M. Pyne, M. Reio, M. Rostes, M. Russo, L. SNEDDON, K. SNow, A. 
Tomaino, and M. Turrty. 2005. Biodiversity values of geographically isolated wetlands: an analysis of 
20 U.S. states. Natureserve, Arlington, VA. 

Eowaarps, A.L.and A.S. Weakiey. 2001. Population biology and management of rare plants in depres- 
sion wetlands of the southeastern coastal plain, U.S.A. Nat. Areas J.21:12-35. 

Feanato, M.L. 1940. A synopsis of Boltonia. Rhodora 42:482-492. 

FerNaco, M.L. 1950. Gary’s manual of botany (ed.8).D.Van Nostrand Co., New York. 

Fiemine,G.P.and N.E.VaN Atstine. 1999. Plant communities and floristic features of sinkhole ponds and 
seepage wetlands in southeastern Augusta County, Virginia. Banisteria 13:67-94. 

Grossman, D.H., D. FaBeR-LANGENDOEN, A.S. WEAKLEY, M. ANDERSON, P. BourGERON, R. CRAWFORD, K. GOODING, S. LANDAAL, 
K. Metzter, K.D. Patterson, M. Pyne, M. Rein, and L. SNeooon. 1998. International classification of ecologi- 
cal communities; terrestrial vegetation of the United States. Volume |. the national vegetation 
classification system: development, status, and applications. The Nature Conservancy, Arlington, 
Virginia. 

Homoya, M. 1983. The upland sinkhole swamps and ponds of Harrison County, Indiana. Proc. Indiana 
Acad. Sci. 92:383-387. 

KaraMAN-Castro, V. and L.E. UrsatscH. 2006. Boltonia. In: Flora of North America Editorial Committee, 
eds. Flora of North America North of Mexico. Vol.20. New York and Oxford. Pp. 353-357. 

Knox, J.S. 1997. A nine year demographic study of Helenium virginicum (Asteraceae), a narrow en- 
demic seasonal wetland plant. J. Torrey Bot. Soc. 124:236-245 

Lassen, K.and W.E. Hamicton. 2003. Competitive interactions between the endangered sink-hole spe- 
cies Helenium virginicum and Boltonia asteroides when exposed to variable water depths. Eco- 
logical Society of America 88"" annual meeting, Savannah, Georgia. (abstract). 

Leonaro, S.W. 1981. Fimobristylis perpusilla Harper in South Carolina. Castanea 46:235-236. 

Leonard, S.W. 1987. Fimbristylis perpusilla in North Carolina. Castanea 52:150. 

McCain, W.E., B.D. McCain, and J.E. Esincer. 1997. Flora of temporary sand ponds in Cass and Mason 
counties, Illinois. Castanea 62:65-73. 

Moraan, J.T. 1966. A taxonomic study of the genus Boltonia (Asteraceae). Ph.D. dissertation, Univer- 
sity of North Carolina, Chapel Hill. 

NatureServe 2005. International ecological classification standard: terrestrial ecological 
classifications.NatureServe Central Databases. Arlington, VA. U.S.A. http://www.natureserve.org/ 
explorer. 

ScHIFFMAN, PM. 1997.Wing reduction in island Coreopsis gigantea achenes. Madrono 44:394-396. 

Smith, M. and T.M. Keevin. 1998. Achene morphology, production and germination, and potential for 
water dispersal in Boltonia decurrens (decurrent false aster), a threatened floodplain species. 
Rhodora 100:69-81. 

Smitu, M. and P Mertier. 2002. The role of the flood pulse in maintaining Bo/tonia decurrens,a fugitive 
plant species of the Illinois River: a case history of a threatened species. In B.A. Middleton, ed. 
Flood pulsing in wetlands: restoring the natural hydrological balance. John Wiley & Sons, Inc., 
New York. Pp. 109-144. 

Snyper, D.B. 2005. List of Endangered Plant Species and Plant Species of Concern. New Jersey Natural 


886 BRIT.ORG/SIDA 22(2) 


Heritage Program, Office of Natural Lands Management, Division of Parks and Forestry, Depart- 
ment of Environmental Protection. 14 pp. 

Stoecker, M.A., M. Smith, and E.D. Mevton. 1995. Survival and aerenchyma development under flooded 
conditions of Boltonia decurrens, a threatened floodplain species and Conyza canadensis,a widely 
distributed competitor. Amer. Midl. Nat. 134:117-126. 

Stevermark, J. 1963. Flora of Missouri. lowa State University Press, Ames. 

Watz, K.S., RJ. Canace, J. Boyte, R. Witte, M.S. Serres, W. HonacHersky, J. Kurtz, and R. Durko. 2001. Identifica- 
tion and protection of reference wetland natural communities in New Jersey: calcareous sink- 
hole ponds of the Kittatinny Valley. New Jersey Department of Environmental Protection, Divi- 
sion of Parks and Forestry, Office of Natural Lands Management, Natural Heritage Program, Trenton, 
New Jersey. 

Weaxtey,A.S.and M.P. ScHarace. 1994. Non-alluvial wetlands of the southern Blue Ridge—diversity in a 
threatened ecosystem. Water, Air, Soil Pollution 77:359-383. 

Wuittacar, R.G., D.F. Durry, and A. Nowrooz!. 1999. Late Cenozoic history of alluvial fans near Stuarts 
Draft, Virginia. Banisteria 13:210. 


oy 


A NEW SPECIES OF WATERLILY 
(NYMPHAEA MINUTA: NYMPHAEACEAE) FROM MADAGASCAR 


Kenneth Landon Richard A. Edwards 
2906 Waco Street 2417 Brass Lantern Road 
San Angelo, Texas 76901, U.S.A. Cedarcreek, Missouri 65627, U.S.A. 


P Ivan Nozaic 


& Veivers Drive 


Sneewah Kuranda 
pb 


Queensland, 4881 AUSTRALIA 


ABSTRACT 


A new species of waterlily, Nymphaea minuta, is described from Madagascar It is closely related to N. stellata 
Willd. but displays a characteristic rompnenon of Sanaa chatacters aud two ewes forms in its life 
re The undersurfaces of the | | to brownish-viol f the taxon. 

mphaea minuta in nature is a dwarf with small eee , cleistogamous flowers. In eulayaiein it exhibits 
two eosin growth forms: a submerged form and an emergent form with floating leaves and larger, emergent, 


chasmogamous flowers. 


Key Worps: Waterlily, Nymphaea, Madagascar 


RESUMEN 
especie de iitto de agua, Nymphaea minuta, de Madagascas, eae Ee emparentada con N. 
ellie Willd. Pero ti én peculiar d y 
ale a Els envés de sus ee es de color gris-pardo a marrén- violace, una caracteristica singular del taxon. 
i aes una planta enana, sumergida En cultivo, muestra 
dos f d imiento diferentes: una ane sumergida y una forma orient et con hojas flotantes y flores 


mas grandes, emergente y casmogaimas. 


The senior authors’ long-time interest and expertise in the cultivation, research, and iden- 
tification of waterlilies has occasioned the present paper. The junior author collected seeds 
from Madagascar that were subsequently grown at the International Waterlily Preserva- 
tion Repository in San Angelo, Texas. Upon examination of these garden-grown plants, it 
became apparent that the material concerned represented an undescribed taxon, the de- 
scription of which follows. 


i des minuta K. Landon, R.A. Edwards & PI. Nozaic, sp. nov. (Figs. 1-4). Type: MADA- 
ASCAR: near village of Tampolo, Coastal Forest in shaded rain pools (ca. 49°...26'E, 17°... 15'S), 1Jul 1999, 
PI. Nozaic s.n. (HOLOTYPE: TEX). 


Nymphaeae stellata Willd. Verd imilis sed differt floribus di libi celal bien 11 | 
et/vel ents portatis et paginis foliorum emergentium supra viridibus infra limoso-canis vel dilute 
brunneolo-violaceis 

Plants rhizomatous growing horizontally, of two distinct forms: submerged and emersed. 
Submerged (underwater) form: leaves very thin, flaccid, ovate, the margins entire and 
wavy, surfaces glabrous, pustulose to wrinkled, bright green above, the veins distinct, lower 
surfaces medium-green with a bronze to reddish cast, the veins bright green, obvious 
and distinct, having 5 primary veins on each side of the midvein, the surface glabrous, 
pustulose to wrinkled, the sinus slightly open with some overlap near attachment to peti- 


SIDA 22(2): 887 — 893. 2006 


888 BRIT.ORG/SIDA 22(2) 


DRAWING 
by PITTMAN 


Fig. 1 Al n ° fA Dian#aith rT 4 1 £] +; | (fi Loa lae \ 


ole; blades mostly 10 cm = 14cm, the petioles usually less in length than the blade, keep- 
ing habit of plant compact and rosette-like in shape; petioles 2-3 mm wide, olive green, 
glabrous. Flowers produced on submerged plants rarely reaching surface, cleistogamous, 
rarely opening underwater; base of cleistogamous bud 0.5-2.0 cm wide, 1.5-2.0 cm long; 
ripe fruit to 3cm = 3cm, petioles slightly curving down as fruit matures, fruit ruptures to 
release seed, seeds brown, oval, smooth, ca. 0.5 mm wide, at release enclosed by aril. Emer- 
gent (above water) form: leaves round, 7-12 cm wide, margins entire to slightly sinuate 
near sinus, sinus open with 10° to 30° angle, the surface medium-green with no other 
pigmentation or marking at any stage of development, lower surface pale violet with 
brown tones, muddy-gray appearance, bright green veins distinctive, petioles olive-green, 
maximum length observed 24 cm, all surfaces free of hairs or scales, overall growth pat- 
tern a circle of ca. 60 cm wide. Flowers small, 2.5-4.0 cm wide; sepals 3 to 4, somewhat 
longer than the petals, tips blunt-pointed, twice the width of petals at the base, sepal 
olive green changing to magenta at base with faint lighter stripes on outside, 7 to 9 veins, 
pale pink to white on inside, with faint greenish stripes. Petals 6 or 7, pale pink to white, 
1.5-2.0 cm long, 0.5 cm at base, tapering to a blunt-pointed tip, with 3 main veins and 2 
smaller veins on the outer edges. Stamens 8-32, shorter than petals, yellow with append- 
ages following petal color. Anthers comprising most of the length, ripe pollen released on 
the Ist day, self-pollinating but also receptive to pollination by other waterlilies in the 
subgenus Brachyceras. Carpels 5 to 15, carpellary appendage a hook-like extension on 
outside wall of carpel, 2-3 mm long. Flowers open near noon and close by late afternoon, 


LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 889 


DRAWINGS 
by PITTMAN 


Fic. 2.N I h inuta. A Seedling B. Bud. C. Sut g {and ial fl D. Aerial flower. E. Fruit 


submerge on third day, open to ca. 45° on first day, and slightly more on 2nd day, often do 
not open on 3rd day, first day flowers usually produce stigmatic fluid, peduncles curve 
down into water as fruit matures. Fruit orbicular, ca. 3 cm x 3 cm, rupturing to release 
seeds; seeds brown, smooth, oval, ca. 0.5 mm wide, enclosed by aril at release. 

Distribution and Ecology.—This species is known in Madagascar only from the type 
locality, where it grows in jungle pools along slow-moving streams. The new species is 
adaptable to growing conditions other than those encountered at the type location and 
easily cultivated, especially in low-light environments. 

Etymology.—The type plants discovered in Madagascar were all dwarf. Under typi- 
cal waterlily cultivation, all plants grown remained dwarf. Only under intensive atypi- 
cal cultivation regimes have larger plants been produced. This species is by far the small- 
est known species of Nymphaea. The epithet ‘minuta’ was proposed by the author who 
discovered this species. After five years of cultivation and multiple generations for con- 
sideration, all authors believe ‘minuta’ to be accurate and appropriate. 


Life history of Nymphaea minuta 

Nymphaea minuta in nature is a dwarf with a submerged habit, including small, cleisto- 
gamous, submerged flowers. These flowers rarely reach the water surface and apparently 
are self-fertilized without the sepals opening or exposing the petals. Various species from 
all subgenera of Nymphaea are self-pollinating, but only N. minuta has been documented 
to do this underwater. Plants of N. minuta in cultivation display both submerged and 


BRIT.ORG/SIDA 22(2) 


s * 


Fic. 3. Top, submerged plants under artificial light in an aquarium, not submerged bud on center plant. Bottom, submerged plant 
grown in pond under nature conditions in a tropical climate. 


LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 891 


disk, | Il | 1 1 I 1 fl fully open. 


892 BRIT.ORG/SIDA 22(2) 


emergent growth forms, the flowers are chasmogamous and vary from floating on the 
water to rising above the surface. Floral morphology and color remain constant in all 
growth forms. In response to changing environmental conditions, an individual plant 
can switch from one form to the other. During these transitions, intermediate phases with 
both floating and submerged leaves are produced, along with both emergent and sub- 
merged flowers (Fig. 2a, 2b). Light intensity and duration appear to be the controlling 
factor in these transitions. 

In temperate latitudes, mature plants of Nymphaea minuta exposed to full sun will 
develop the emergent phase throughout the summer but revert to the submerged phase 
as the daylight lessens in length and intensity. The same plant often reverts back to the 
emergent phase the following summer. 

Almost all cultivated specimens have been dwarf. However, under cultivation with 
fertilization and optimum conditions of light and climate, plants have been produced 
roughly four times larger (including flowers and all vegetative parts of the plants) than 
the dwarf plants first discovered. Seeds [of cleistogamous and chasmogamous flowers] 
remain the same size, although the number of seeds produced from the larger flowers 
and resulting larger fruits, increases proportionally to the rest of the plant. The number 
of carpels and stamen also increase at the same ratio as the size increase of vegetative 
parts. 

Seedlings produced in full sun in late spring through mid summer will immediately 
grow into an emergent plant, with floating leaves and flowers at or above the water sur- 
face, typical of brachyceras tropicals. Seeds germinated in the late summer or fall will 
often grow in the submerged form until the following summer or until artificial light of 
enough intensity and duration are provided, causing the plant to transform into the emer- 
gent form. Seedlings grown in shade remain in the submerged phase but continue to grow 
and produce flowers and fertile fruits. 

In chasmogamous flowers, stigmatic fluid is usually produced on the first day of 
anthesis, and ripe pollen is released. As the fruit ripens, the peduncles bend downward 
and fruits mature beneath the water surface. 

As the plants mature, a perennial rhizome is produced. Dormant tubers have been 
produced in cultivation and would be expected to occur in the wild populations in ephem- 
eral conditions. 


Relationship of Nymphaea minuta to other species 

Characters of the flowers place the new species in Nymphaea subgenus Brachyceras. The 
cary ee at the sides with separate walls for each ovary. Carpellary styles are present 
asa small hook-like extension at the back of each carpel. The stamens have distinct ap- 
pendages beyond the anthers. 

Nymphaea minuta at a glance ee more closely resembles Nymphaea 
stellata Willd. (not N. nouchali Burm. f., a different species, in our opinion) of tropical 
Asia than any other known species of aoe Both species display a coarsely dentate 
margin on floating leaves. Flowers display very low petal numbers in both species. The 
type habitat for N. minuta would also support N. stellata although ina stunted form. Such 
plants would in fact more closely mimic the dwarf nature of N. minuta rather than a 
typically proportioned N. stellata. 

Upon closer inspection several differences between the two species would become 
apparent. Nymphaea minuta has no contrasting pigmentation upon the leaves in either 


LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 893 


form. Nymphaea stellata often display slight mottling or flecks of purple pigmentation 
on floating leaves as they mature. The lower surface of the leaves of both species exhibit 
distinct and different venation and pigmentation patterns. Like most Nymphaea species, 
N. stellata produces submerged leaves as seedlings and emerging from dormancy, but 
they are diminutive, quickly deteriorate and do not represent a submerged growth form. 
Nymphaea minuta produces substantial and persistent submerged foliage, flowers and 
fruits, completing a full life cycle completely submerged. 

Nymphaea minuta apparently will hybridize with other species and cultivars of 
subgenus Brachyceras. With N. minuta as pollen donor, cross-pollination was attempted 
on N. micrantha Guill. & Perr. and N. colorata Peter. and seed development proceeded in 
normal fashion. Similar results were obtained using three additional cultivars as seed 
parents. Nymphaea minuta was also used as the seed parent with crosses to hybrids with 
distinctive features, such as pigmentation on leaves. Further tests must be performed to 
see if cross-pollination actually occurred (vs. asexual seed development), but both senior 
authors have progeny from crosses displaying obvious hybridization. Crosses within the 
subgenus Brachyceras, with resulting progeny displaying obvious characteristics, sucha 
pigmentation on the leaves similar to the pollen parent, provides further evidence this 
species should be placed within the subgenus Brachyceras. 

Dimorphism in growth form is a rare feature in the genus Nymphaea. No published 
material exists discussin g this feature with any other known subgenus of Nymphaea. At 
the International Waterlily Preservation Repository, we have taxon in the subge- 
nus Brachyceras that appears to exhibit some degree of persistent submerged growth, 
however, the research is incomplete. 


ACKNOWLEDGMENTS 
We are grateful to Guy Nesom for the Latin diagnosis and for a detailed review of the 
manuscript and to B.L. Turner for his encouragement and help with this project. We also 
thank Ed Schneider for a thorough review of the paper. Drawings for the new species 
were kindly provided by Katherine Pittman. 


REFERENCES 


Conard, H.S., 1905. The waterlilies: a monograph of the genus Nymphaea. Publ. Carnegie Inst. Wash- 
ington. No. 4. Carnegie Institution Washington, Washington, DC. 


BRIT.ORG/SIDA 22(2) 


Book REVIEW 


Donato G. Crossy. 2004. The Poisoned Weed, Plants Toxic to Skin (ISBN 0-19-515548-3, 
hbk.). Oxford University Press. (Orders: Oxford University Press, 2001 Evans Road, 
Cary, NC 27513, U.S.A., 800-451-7556, 919-677-1303 fax, wwwoup.com). $59.95, 214 


pp. figures, graphs, 24 color figures, 6 1/2" x 91/2" 
about the plants that 


Dermatoxic plants are all around us. The book entitled helps readers learn much more a 
cause itchiness, rashes and other very painful symptoms. This book gives a detailed background with compre- 
chemicals involved in causing dermatoxic reactions. 

first gives a background history of a variety of plants that cause 


Informatior n the 


hensive, technical 

The book consists of ten chapters. The 
reactions such as allergic contact dermatitis (ACD) in humans. Much of this history centers on the naming, his- 
ived causes of the ACD reaction from poison ivy and its relatives. The second chapter focuses on 
ir relatives. The third chapter discusses other 


—_ 


tory and 

the taxonomy, habitat and distribution of the poison ivies and the 
plants that can cause ASP in humans. Plants covered by the “other allergens” include wild species such as 
onwood), Quinone en. plants such as the amano 20 trees or ex- 


id 
\ 
recian lau- 


fetopium 


otic bardaroads — eacel (Dalbe is gid sp.), | lactone containing § species such as Lat 


ts and others 


rel), weeds, food pl 
chapter four discusses preetone pes from the Rutaceae family, Apiaceae family, and Moraceae nae 
and Urticaceae families. Chapter five 


| Brassicaceae, Alliacea 


Fats 6a 


along with irritant plants from 
is a chapter detailing chemical and physical properties of the allergens related to urushiol. This nee ation 


meuniees aan ication ane anal has ondation, puoi. and bio-degradation, reactions with proteins and more. 
[ | properties of allergens such as quinines, lactones, acety- 
sand rubber latex. Chapter seven discusses the physical and chemi- 


(oni alcohols, essential oils, lic] b 
cal properties of phyioreac and irritant (non- a ae ee neuen Furocoumarins, Hypericins, Irri- 
xalates, Chapters eight and nine 


— 


tant esters, organosulfut ( ompounds 
— forms of exposure to various ACD substances and their adverse effects. 

hapter ten includes methodologies to prevent exposure to ACD substances as well as discussion on treat- 
ment options. The chapters are followed by multiple appendices, containing chemical structures and physical 
Calis for various substances discussed in the text and some key references. Indices are separated into plant 


a general index 


inder common names and 
The Poisoned Weed: Plants Toxic to Skinis a very aa mative and timely book. P rimarily designed for pro- 
elds, it looks to be the new authority on dermatoxic 


name 


fessionals in the biochemical, chemical, and medicina 
plants and substances which cause allergic contact pees Luckeydoo, Herbarium, Boe Research 


Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A 


SIDA 22(2): 894. 2006 


TWO NEW COMBINATIONS IN PEYRITSCHIA 
(POACEAE: POOIDEAE: AVENINAE) 


Victor L. Finot Paul M. Peterson 
eae de Concepcion Department of Botany 
nomia eee ee 
Depart to de Produccién Animal mithsonian fe 
Casilla 537, Chilldn, CHILE ie DC 20013 ae U.S.A, 
vifinot@udec.cl ae, 


Fernando O, Zuloaga 
Instituto de Botanica Darwinion 
3 0 
San Isidro 1642, ARGENTINA 
fzuloaga@darwin.edu.ar 
ABSTRACT 


Trisetum howellii Hitchc., an endemic species from the Galapagos Islands, Ecuador, and Trisetum pinetorum 
wa Ae an aos Species pom Guatemala, are pineal into Peyritschia IP. howell fEitehe. ) Finot & PM. 


PM. Peterson]. P 


with new areas of distribution documented in South America. A key disti he species and able com- 
paring the salient morphological features of all species within be are erouded 


RESUMEN 


Tri howellii Hitchc., especie endémica de | la cuador, y Trisetum pinetorum Swallen, especie 

endémica de Guatemala se transfieren a Peyritschia [P. howellii ieee Finot & P.M. Peterson; P. pinetorum 

(Swallen) Finot @& P.M. Peterson]. Peyritschia, asi circunscrita, incluye siete especies con nuevas areas de 

distribucion documentades en Sudan ence) Pioporeona una epi peas distinguir las especies y un cuadro 
onde se compar g las especies de Peyritschia. 


The genus Peyritschia was described by Fournier (1886) and originally included only 
one species, P. koelerioides (Peyr.) E. Fourn., based on Aira koelerioides Peyr, from south- 
ern México and Guatemala (Finot et al. 2004). A second species from México, Deschampsia 
pringlei Scribn., was transferred to Peyritschia by S.D. Koch in 1979[= P. pringlei (Scribn.) 
S.D. Koch], and is also known from Guatemala, Costa Rica, Venezuela, and Ecuador 
(Hernandez-Torres & Koch 1988; Pohl & Davidse 1994; Finot et al. 2004). This small ge- 
nus (Peyritschia) was later included within Trisetum by Hernandez-Torres and Koch 
(1987). When placed in Trisetum, P. koelerioides and P. pringlei are named: Trisetum 
altijugum CE. Fourn.) Scribn. and T. kochianum Hern.-Torres., respectively. 

Peyritschia has isomorphic to sub-isomorphic, linear, 1-3-nerved glumes; bilobed 
lemmas that are awned near the base or the middle of the back or the awn reduced toa 
subapical mucro; paleas that are tightly enclosed by the margins of the lemma; linear 
lodicules; and an androecium composed of two stamens. In contrast, Trisetum has het- 
eromorphic, ovate-lanceolate or oblanceolate, 1-5-nerved glumes |first glume 1(-3)-nerved, 
second glume 3(-5)-nerved|]; lemmas with 2 to 4 short setae at the apex due to the prolon- 
gation of the nerves and a dorsal awn born on the upper third of the lemma, rarely lo- 
cated near the middle of the back; paleas that are free from the margins of the lemma; 
lodicules with 2- or 3-lobed apices; and an androecium with three stamens (Finot et al. 


SIDA 22(2): 895 — 903. 2006 


896 BRIT.ORG/SIDA 22(2) 


2004, 2005a, 2005b). Peyritschia includes perennial herbs with flat leaf blades, mem- 
branous ligules, narrow to contracted-spiciform or lax and somewhat open panicles, 2(- 
3)-flowered spikelets, rachillas disarticulating above the glumes and between the florets, 
glabrous ovaries, and caryopses with liquid endosperm. Previous studies showed that the 
micromorphology of the lemmatal epidermis has good characters to distinguish Trisetum 
from Peyritschia. Most species of Peyritschia lack prickles hairs (present in all species of 
Trisetum), although, if present, they are restricted to the keel or the apex of the lemma in 
P. deyeuxioides and P pinetorum. All species of Peyritschia have bordered hooks alter- 
nating with epidermal long cells (hooks not bordered, nor alternating with long cells in 
Trisetum) and lack macrohairs (macrohairs are present in some species of Trisetum)[Finot 
et al. 2006]. These characteristics along with the salient features of the spikelet, were con- 
sidered important for resurrecting the genus by Finot (2003a) who transferred three ad- 
ditional species into Peyritschia: P. conferta (Pilg.) Finot (=Trisetum confertum Pilg.) from 
Bolivia, Ecuador and Venezuela; P deyeuxioides (Kunth) Finot (=T. deyeuxioides Kunth) 
widely distributed in México, and extending into Central and South America (Colombia, 
Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Nicaragua, Panama and Venezu- 
ela); and P. humilis (Louis-Marie) Finot (=T. humile Louis-Marie), an endemic in México 
(Finot et al. 2004). 

Trisetum confertum was originally described from an Ecuadorian specimen (Crescit 
in Prov. Imbabura, ad Loma de Canaballa et locis vicinis; alt. 2100-2300 m; A. Sttibel 152: 
holotype, B; isotypes, US-81771 ex B!, CONC fragm. and photo ex B!). Hitchcock (1927) 
cited this species as occurring in Cochabamba, Bolivia. Later, Valencia (1941) moved the 
species to Deschampsia[D. conferta (Pilg.) Valencia]. However, Parodi (1949) and Chiapella 
(2000) excluded it from Deschampsia. 

More recently, two species, Irisetum de yeuxioidesand T. humilis, were transferred in 
Peyritschia, based on spikelet and floret characteristics (Finot 2003a; Finot et al. 2004). 

Three of the five currently recognized aps o oe il Sot id are found in Guatemala: 
P. koelerioides (the type species of the g thern México; P. pringlei 
also present in México, Costa Rica, Venezuela, and Ecuador; and P. deyeuxioides ranging 
from southern México to Ecuador (Pohl & Davidse 1994; Espejo-Serna et al. 2000; Finot 
2003a; Finot et al. 2004). 

The following seven species of Trisetum have been mentioned as occurring in Guate- 
mala: I’ angustum Swallen from San Marcos near Volcan Tajumulco, also extending into 
southern México; T. pinetorum Swallen here transferred to Peyritschia, from 
Quetzaltenango near Volcan Santo Tomas, and now known from México; T. viride (Kunth) 
Kunth (syn. T. altum Swallen) from El Progreso also widespread in México; T. irazuense 
(Kuntze) Hitche. from Huehuetenango near Tunima, mentioned for the first time by Finot 
et al. (2004), also from Costa Rica, Honduras, Panama, Colombia, Venezuela, Ecuador, 
and Peru; I. rosei Merr. from Huehuetenango from near Volcan Santa Maria, also extend- 
ing into México; T. pringlei (Scribn. ex Beal) Hitchc. from Huehuetenango and 
Totonicapan, also found in México, Costa Rica, and Panama; and T. spicatum (L.) K. Richt. 
with a wide distribution in the Americas (Swallen 1955; Pohl 1980; McVaugh 1983; 
Hernandez-Torres 1988; Pohl & Davidse 1994; Finot 2003b, Finot et al. 2004, 2005a, 
2005b). 

In Ecuador, the following four species of Trisetum have been reported: T. andinum 
Benth., endemic to Ecuador; T. spicatum and T. oreophilum Louis-Marie var. oreophilum 
ranging from Ecuador to Pert and Bolivia (Finot et al. 2005b); and T. howellii here trans- 


FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 897 


ferred to Peyritschia. Species of Peyritschia found in Ecuador are: P deyeuxioides and P 
pringlei (Finot 2003a). 

Preliminar Vi | using Mor phological characters within Trisetum 
s.l. (including Trisetum. s.s., Peyritschia, and Sphenopholis) [Finot 2004], depicts a well- 
supported clad (bootstrap of 95 %) that includes Peyritschia koelerioides, P pringlei, P 
deyeuxioides, P. conferta, and T. howellii Hitchc. This clade is supported by three syna- 
pomorphies: terete lemmas in lateral view, two stamens, and ornamentation of the lemma. 

In this paper we make two new combinations, Peyritschia pinetorum(Swallen) Finot 
& PM. Peterson, from Volcans Atitlan and Santo Tomas, Guatemala, and P howellii 
(Hitchce.) Finot & PM. Peterson, from Galapagos, Ecuador. In addition, we include a key to 
distinguish all seven species, a table comparing their salient morphological features, and 
a new illustration, description, and SEM photo of the lemmatal surface of P. pinetorum. 


Peyritschia howellii (Hitchc.) Finot & PM. Peterson, comb. nov. (Table 1). Trisetum howellii 
Hitchc., Proc. Calif. Acad. Sci. ser. 4, 21(24):296. 1935. Type: ECUADOR. GALAPAGOS ISLANDS: Indefatigable 
Island, Mt. Crocker, 9 May 1932, J.T. Howell 9208 (HOLOTYPE: CAS-211262!: ISOTYPE: US-1611545)). 


Perennials; culms 35-60 cm tall, weak; internodes glabrous, 5-6 nodes per culm, nodes 
glabrous. Leaf sheaths shorter than the internodes, glabrous, striate; ligules 0.5-1.5(-2) 
mim long, apex obtuse, not ciliate ti denticulate, glabrous; blades 8-15cm x 2mm, 
flat, glabrous, smooth, margins sometimes scabrous. Panicles 8-18 cm long, (0.5-)10-20 
mm wide, contracted, narrow, lax, somewhat open, exserted or sub-included in the up- 
per sheath; rachis and pedicels glabrous; spikelets 4.5-5.5 mm long, (1-)3-flowered; 
rachillas 1.5 mm long, copiously pilose, the hairs ca. 1.5 mm long, as long as the rachilla: 
glumes almost covering the florets, isomorphic, linear to linear-lanceolate, narrow, some- 
what scabrous on the keel, the margins hyaline, apex acute, purplish; first glumes 4.3-4.7 
x 0.3mm, l-nerved; second glumes 4.7-5.5 x 0.4 mm, 1-3-nerved; first lemmas 4-4.5 = 0.5 
mm, linear-cylindrical to linear-lanceolate, glabrous, smooth, or scaberulous only towards 
the apex, purple towards the apex, rounded on the back, the margins enclosing the palea, 
5-nerved, awned, the awn 5.5-7 mm long, twisted, geniculate, somewhat scabrous, borne 
on the upper third of the lemma, at 1-1.5 mm below the apex, the apex bidentate, toothed, 
acute with the intermediate and marginal nerves conspicuous towards the apex; callus 
obtuse, with stiff hairs, the hairs 0.5 mm long; paleas 3-4.3 mm long, shorter than the 
lemmas, 2-nerved, the nerves scabrous owas the apex, apex acute, ale ovary gla- 
brous; lodicules 0.6 mm long, linear, entire; stamens 2, anthers about 0.8 mm long. Cary- 
opses 2 mm long, glabrous; endosperm liquid. 

Phenology.—Flowering from September to January. Matures fruits are found from 
February to March. 

Distribution and habitat.—Peyritschia howellii is an endemic species from Ecuador. 
Most of the studied collections come from Isla Santa Cruz (Indefatigable), Galapagos, grow- 
ing between 480-1000 m; Jeppesen 76 (AAU) is the first collection of the species made 
from Isla San Cristobal (Chatham). 

Comments.—Peyritschia howellii was included by Valencia et al. (2000) in the Red 
Book of the endemic plants of Ecuador (as Trisetum howellii). 


Additional ined. ECUADOR. Galapagos Islands: Isla Santa Cruz, in area that includes W slope of 

Media Luna, E slope of Cerro de los Caminos and flat plain between, 4 Sep 1975, D.A. & D.B. Clark 447 (AAU); 
Puntudo, 17 Mar 1977, A. & H. Adsersen 1413 (QC - ca. uN m, 8 Nov 1966, U. & I. Eliasson 557 (S); sur de Mt. 
Crocker, 710 m,17 Oct 1966, U.& I. Eliasson 284(S) Bella Vista, 480 m, 19 Oct 1966, U. & I. Eliasson 356 (S); 
no date, Fagerlind & Wibon 2957 (S); no date, Paver lind & Wibon 3320(S); highest mountain top, 860 m, 23 May 


868 


Taste 1. Salient morphological characteristics to distinguish among the seven species of Peyritschia. 
P.conferta P. deyeuxioides P. howellii P. humilis P. pinetorum P. koelerioides P. pringlei 
Panicle shape narrow, contracted — lax, somewhat narrow, narrow, contracted  narrow,contracted — spiciform narrow, 
open and nodding somewhat open somewhat open 
Number of florets 2 2073 (1-)3 2 2 2or3 
per spikelet 
Spikelet, length (mm) 5.0-55 4.5-8.0 4.5-5.5 -7 - 5 
Glumes, shape oval-lanceolate linear linear to linear- lanceolate lanceolate to lanceolate to lanceolate 
lanceolate ovate-lanceolate ovate-lanceolate 
Glumes, size verses longer equal, shorter or equal longer slightly shorter longer as long as the 
spikelet longer florets or longer 
Glumes, length (mm) 5.2-5.5 (3.5-)4-5.5 (-8) 4.3-5.5 4 45-6 3-5 
Lemma, length (mm) 3.5-5.0 4-6 4-45 3-3.5 5-6 2.5-4 3-43 
(lower floret) 
Lemma, apex bilobed, lobes bidentate, toothed, bilobed, lobes bifid, toothed, bilobed, lobes bilobed, lobes 


obtuse 
Rachilla,indument sparsely pilose 


Awn, position basal 

Awn, shape geniculate, twisted 
Awn, length (mm) 6-7 

Callus indument short pubescent 


entire or bilobed to 
bidentate 
copiously pilose 


median 
geniculate, twisted 


4.5-8(-12) 


short pubescent 


acute 

copiously pilose 
upper third 
geniculate, twisted 


short pilose 


obtuse 

sparsely pilose 
middle or lower 
third 

geniculate, twisted 


sparsely pilose 


acute 

copiously pilose 
pubescent 
median 


geniculate, twisted 
7-12 
pilose 


btuse 
glabrous to sparsely 


apical if present 
(as a short mucro) 
straight or absent 


glabrous to short 
pubescent 


obtuse 
sparsely pilose 


basal 


geniculate, twisted 


short pubescent 


(Z)% vals/OYO'LINA 


FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 899 


1959, Harling 5764 (S), 1850 ft, 27 Jul 1974, van der Werff 1330 (S); Mt. Crocker, 840 m, 18 Feb 1967, Wiggins & 
Porter 652(S); fern-sedge zone along trail to Mt. Crocker, 500 m, 6 Feb 1964, Wiggins 18577 (S); Isla San Cristobal, 
El Junco, 650 m, 28 Feb 1976, Jeppesen 76 (AAU). 


aa L Septet (Swallen) Finot & P.M. Peterson, comb. nov. (Fig. 1, Table 1). Trisetum 
um Swallen, Phytologia 4:424. 1953. Type: GUATEMALA. QUEZALTENANGO: Volcan Santo Tom 
on pine- ines clad slope, 2500-3700 m, 22 Jan 1940, ].A. Steyermark 34824 (HOLOTYPE: F-1048257! ISOTYPES: 
MO fragm. ex FI, US-2240525 ex Fl, US-2236478 fragm. ex F)). 


Perennials; culms 33-70 cm tall, weak, internodes glabrous, 3-4 nodes per culm, nodes 
glabrous. Leaf sheaths shorter than the internodes, glabrous, striate; ligules 1.5-3 mm long, 
truncate, dentate-laciniate, glabrous; blades 5-10 cm long, 1-1.5(-2) mm wide, flat, gla- 
brous above and below. Panicles 5-15 cm long, 5-10 mm wide, contracted, narrow, some- 
what eS somewhat interrupted, exserted or sub-included in the upper leaf sheath; ra- 
chis g] Is scabrous or glabrous. Spikelets 5-7 mm long, 2-flowered; rachillas 
ca. 1 mm lene. copiously pilose, the Haus about 1-2 mm long; glumes slightly shorter 
than the florets, sub-isomor phic anceolate, the keel smooth to slightly 
scabrous towards the apex, apex acute; first glumes 4.5-6 x 0.7-0.8 mm, a little shorter 
and narrower than the second glumes, l-nerved; second glumes 5-6 x 0.7-0.8 mm, l-or 3- 
nerved; first lemma about 5-6 mm long, lemmas glabrous, awned, rounded on the back 
with involute margins on the lower 1/2 enclosing the palea, the awn 7-12 mm long, twisted, 
geniculate, and inserted on the median portion of the back, the apex deeply bifid with 
acute teeth, each tooth with two short awns derived from the prolongation of the mar- 
ginal and intermediate nerves; callus obtuse, with short hairs, the hairs ca. 1 mm long; 
paleas about 3.5-4 mm long, shorter than the lemmas, 2-nerved, the nerves scabrous to- 
wards the apex, apex acute; lodicules about 0.5 mm long, linear, sometimes with a little 
lateral lobe near the median portion, the apex subacute; stamens 2, anthers 0.9-1.5 mm 
long; eel glabrous. Caryopses not seen. 

mma micromorphology. Long cells rectangular, 3-12 times longer than wide; anti- 
ae ents parallel and highly undulate, the undulations U- or V-shaped; periclinal walls 
flat; silica cells about 10 um in diameter, short, elliptical; stomata absent; prickle hairs 
about +5 um long, ovate-elliptical, restricted to the keel and with a short apical barb; 
long cells about 10-12.5 um in diameter, alternating with bordered hooks; macrohairs 
absent (Fig. 2.). 

Phenology.—Flowering in January. 

Distribution and habitat.—Known only from southwesterm Guatemala in Departa- 
mentos Quetzaltenango and Solola. Apparently, P pinetorum is restricted to volcanic soils 
since the only two localities are from higher elevations (2500-3700 m) on slopes of Volcan 
Atitlan and Santo Tomas. 

Comments.—This new combination can be distinguished from all other known spe- 
cies of Peyritschia by having lemma apices that are deeply bifid, and these apices end in 
four hyaline setae derived from the apical extension of the marginal and intermediate 
nerves (Table 1). The presence of these setae suggests that P. pinetorum, ina phylogentic 
sense, lies very near Trisetum. Nevertheless, because of the shape of the lodicules (entire 
in Peyritschia and bilobed or toothed in Trisetum), the lemmas rounded on the back (ver- 
sus keeled in Trisetum), the awn inserted on the median portion of the lemma [inserted 
on upper 1/3 in Trisetum except in Trisetum subgen. Deschampsioideum (Louis-Marie) 
Finot], paleas that are tightly enclosed by the margins of the lemma (gaping in Trisetum 


— 


ite 
eee. 


oe 
KK: 


eX 


SSK 
‘es 
a 


A. Habit. B. Sheath 


liaule 
y 
eee 


d 
t 


BRIT.ORG/SIDA 22(2) 


f the blade. C. Spikelet. D. First floret. E. Palea, dorsal view.F. Palea 
F 4 Y 


Fic. 1. Pe 


is, dorsal view. J. 0 
Lf é T és é F 


is, ventral view 


J b 
ventral view. G. Rachilla. H. Lodicules.1.0 


P 


FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 901 


# 


Fic ») CEM nhat bh af tho | | ef ANN\ .£ D reps Pe) H rr {C+ L 2AQDVA\ ch 
2 / \ 


nating with long cells. Scale bar = 55m. 


and not tightly enclosed by the margins), an androecium with two stamens (3 stamens in 
Trisetum), and lemmatal epidermis with bordered hooks alternating with long cells 
(verses epidermis without bordered hooks, the hooks not alternating with long cells in 
Trisetum) [Fig. 2.], we place the new species in the genus icibeciaee 


[Or 


i 
5 
o) 
Sy 
& 
a 
a 
pe 
am 
aN 
wr 
e; 
Gi 
oO 
Co 
Z, 
Oo 
ha 
3 
No) 
ron 
ee 
= 
=> 
a 
wi 
io) 


Additional specimen examined. GUATEMALA. Solola: Volcan Atitl: 
(3540 m), 23 Jan 1907, W.A. Kellerman s.n. (US-2181368). 


KEY TO THE SPECIES OF PEYRITSCHIA 


. Lemma without a dorsal awn, apex muticous or with a subapical mucro P. koelerioides 
. Lemma with a dorsal awn, the awn geniculate and extended beyond the glumes 
2. Rachilla copiously pilose, the hairs 1-3 mm long; awn borne on the middle or the upper 

third of the back of the lemma 

3. Lemma with the apex deeply bifid; panicles 5-15 cm long, contracted, narrow 


P. pinetorum 


902 BRIT.ORG/SIDA 22(2) 


3, Lemma with the apex entire or shortly bidentate or bilobate; panicles 8-35 cm long, lax, 
somewhat open 
4. Rachillas with hairs 2-3 mm long; glumes linear 5.2-5.5 mm long; awn borne on the 


middle portion of the lemma P. deyeuxioides 
4. Rachillas with hairs about 1.5 mm long; glumes linear to linear-lanceolate 4.3-5.5 mm 
long; awn borne on the upper third of the lemma P. howellii 


2. Rachilla pubescent, the hairs 0.2-0.8 mm long; awn borne near the base of the lemma. 
5. Culms 5-12 cm tall; leaf blades 2.3-3.5 cm long; panicles about 2.5 cm long, 0.5 cm wide 
P. humilis 


5. Culms 20-200 cm tall;leaf blades 5-15 cm long; panicles 5-20 cm long, 1-4(-4) cm wide. 

6. Spikelets 2- or 3-flowered, 4-5.5 mm long; glumes 4-5.3 mm long; lemma 3-4.3 mm 
long P. pringlei 

6. Spikelets 2-flowered, 5—5.5 mm long; glumes 5.2-5.5 mm long; lemma 3.5-5 mm long 
P.conferta 


ACKNOWLEDGMENTS 


We thank the Directors and Curators of the following herbaria: AAU, BA, BAA, BAF, C, 
CR, CONC, F LP, MERL, P PR, QCA, SGO, §S, SI, US. The first author gratefully acknowl- 
edges the Myndel Botanical Foundation for a fellowship to study types of Trisetum and 
allied genera at Paris (P) and Stockholm (S); a grant from MECESUP-Universidad de 
Concepcion UCO-9906/01 to study Trisetum at the Smithsonian Institution (US) in Wash- 
ington, DC, U.S.A and Instituto de Botanica, Darwinion (SI) in Buenos Aires, Argentina; 
Projects DIUC 204.121.009-L0 and 205.111.047-1.0 for financial support; Oscar Matthei 
and Clodomiro Marticorena for directing my Ph.D. studies, and Susan J. Pennington (US) 
for helping me during my stay at the Smithsonian. In addition we thank Vladimiro Dudas 
(SD for the beautiful illustration and Yolanda Herrera Arrieta and Ana Maria Planchuelo 
for reviewing the manuscript. This paper is part of the doctoral thesis of the first author 
at Departamento de Botanica, Universidad de Concepcion, Concepcion, Chile. 


REFERENCES 

Cuiapetta, J. 2000. Taxonomy, morphology, phylogeny and biogeography of Deschampsia (Poaceae- 
Aveneae). Universidad San Carlos de Bariloche, Doctoral thesis. 

Espeso-Serna, A., A.R. Lopez-Ferrari, and J. Vacpes-Reyna. 2000. Poaceae. In: A. Espejo-Serna and A.R.L6pez- 
Ferrari (eds.). Las Monocotiledéneas Mexicanas. Poaceae Barnhart. Una Sinopsis Floristica. Partes 
IX—XI. Pp. 8-236. 

Finor,V.L. 2003a. Peyritschia.P.478.In: Soreng, R.J.,PM. Peterson, G. Davidse, EJ. Judziewicz, F.O.Zuloaga 
TS. Filgueiras, and O. Morrone. 2003. Catalogue of New World grasses (Poaceae): IV. subfamily 
Pooideae. Contr. U.S. Natl. Herb. 48:1—730. 

Finor, V.L. 2003b. Trisetum. In: Soreng, R.J., PM. Peterson, G. Davidse, E.J. Judziewicz, F.O. Zuloaga, TS. 
Filqueiras,and O.Morrone. 2003.Catalogue of New World grasses (Poaceae): IV. subfamily Pooideae. 
Contr. U.S. Natl. Herb. 48:659-676. 

Finot, V.L. 2004. Sistematica del género Trisetum Pers. (Poaceae: Aveneae) en América. Una 
aproximacidn filogenética basada en datos morfoldgicos. Universidad de Concepcion, Doctoral 


thesis. 

Finor, V.L., PM. Peterson, R.J. Sorenc, and F. Zutoaca. 2004. A revision of Trisetum, Peyritschia, and 
Sphenopholis (Poaceae: Pooideae: Aveninae) in México and Central America. Ann. Missouri Bot. 
Gard. 91:1—30. 

Finor, V.L., PM. Peterson, R.J. Sorene, and F.O. ZuLoaca. 2005a. A Revision of Trisetum and Graphephorum 
(Poaceae: Pooideae: Aveninae) in North America. Sida 21:1419-1453. 


FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 903 


Finot, V.L., RM. Peterson, F.O. ZULOAGA, R.J. SoreNG, and O. MattHel. 2005b. A Revision of Trisetum (Poaceae: 
Pooideae: Aveninae) in South America. Ann. Missouri Bot. Gard. 92:533-568. 

Fino, V.L., C.M. Baeza, and O. MattHel. 2006. Micromorfologia de la epidermis de la lemma de Trisetum 
y géneros afines (Poaceae: Pooideae). Darwiniana.|n press. 

Fournier, E. 1886. Mexicanas plantas Il. Gramineae 1-xix, 1-160, pl. 1-10. Paris. 

HerRNANDEZ-Torees, |. and S.D. Koch. 1987. The status of the genus Peyritschia (Gramineae: Pooideae). 
Phytologia 61:453-455, 

HerNANnbez- Torres, !.and S.D. Kock. 1988.Revisidn taxondmica del género Trisetum (Gramineae: Pooideae) 


en México. Agrociencia 71:71-102. 

HitcHcock, A.S. 1927. The grasses of Ecuador, Pert, and Bolivia. Contr. U.S. Natl. Herb. 24:291-556. 

HitcHcock, A.S. 1935.The Templeton Crocker expedition of the California Academy of Sciences, 1932, 
No. 24,new species of grasses from the Galapagos and the Revillagigedo Islands. Proc. Calif. Acad. 
Nat. Sci.,ser.4 21:25-300. 

Koch, S.D. 1979. The relationships of three Mexican Aveneae and some new characters for distin- 
guishing Deschampsia and Trisetum (Gramineae). Taxon 28: 225-235. 

McVauch, R. 1983. Gramineae. Flora Novo-Galiciana 14:1-436. 

Parooi, L.R. 1949. Las gramineas sudamericanas del género Deschampsia. Darwiniana 8:415-475. 

PoHt. RW. 1980. Gramineae. In:W. Burger, ed. Flora Costaricensis. Fieldiana Bot. n.s. 4:1-608. 

Pout, R.W.and G.Daviose. 1994. Trisetum Pers. ln:G. Davidse, M. Sousa, and A.O.Chater (eds). Fl. Mesoamer. 
G253=23 3. 

Swaiten, J.R. 1955. Gramineae. In: PC. Standley and J.A. Steyermark, eds. Flora of Guatemala, Part Il: 
Grasses of Guatemala. Fieldiana Bot. 24:1-390 

VALENCIA, J.1. 1941. Especies criticas de Trisetum que deben pasar al género Deschampsia. Revista Ar- 
gent. Agron. 8:1 22-130. 

VALENCIA, R., N. PITMAN, S. LEON-YANEZ, and P.M. Jorcensen (eds.). 2000. Libro Rojo de las Plantas Endémicas 
del Ecuador 2000. Pontificia Universidad Catolica del Ecuador, Quito. 


904 BRIT.ORG/SIDA 22(2) 


Book REVIEW 


Marr Waite. 2000. Prairie Time: A Blackland Portrait. (ISBN 1-58544-501-0, hbk.). Texas 
A&M University Press, 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 
979-458-3982, 979-847-8752 fax; http://www.tamu.edu/upress/). $19.95, 272 pp., 
b/w figures, 1 map, index, 51/2" x 81/2" 

Texas-prairie enthusiasts will relish author Matt White’s new memoir of the Blackland Prairie called Prairie 


Time: A Blackland Portrait. His new book intertwines the author's interests in the Blackland Prairie with per- 


fek 


sonal touches of family, natura, and land histories, with prairie ecology—all with the background of his senti- 
mental memories of the Blackland Prairie region. 


The book is a collection of Matt White's experiences in exploring prairie remnants; among these are his 


ay 


interactions with land stewards and also stories of unfortunate losses of prairie remnants to various causes. 
Throughout the book there is information on animals that have inhabited the prairie, such as the prairie chicken 


and wild turkeys, as wel 


as information on the importance of prairie as wildlife habitat. Prairie ecology is dotted 
through the text as well. For eXé ae, yowll find information on the importance of fire to the prairie system, the 
for t] | ch . t 


stics of prairie soils, flora indicative of prairie areas, and flora invasive to the prairie. A 
discussion of historical land uses and views of the prairie over time is also included. The author's deep interest 
and enjoyment of the prairie is delicately sprinkled throughout the text. 

Take a sentimental journey with author Matt White and experience his excitement in a aa ing I ee 
ad | 


Prairie remnants in Texas. White shares both joyous experiences of preserving remnants an 
feats in loss of Blackland Prairie areas. Readers with an interest in learning more about the Blackland Paine 


should seek out this very readable book—it includes many details on the natural history and ecology of the Black- 


land Prairie within the author's stories. es Luckeydoo, Herbarium, Berane al Research iia of Texas, 509 
Pecan Street, Fort Worth, 1X 76102-4060, U.S. 


SIDA 22(2): 904. 2006 


POA MATRI-OCCIDENTALIS (POACEAE: POOIDEAE: 
POEAE: POINAE), A NEW SPECIES FROM MEXICO 


Paul M. Peterson and Robert J. Soreng Yolanda Herrera Arrieta 
Department of Botany CHDIR- Dgo, ae 
ee Wen im elinee History Instituto Polit 
nstitution Sigma s.n. Fracc. 20 de Noviembre II 
ee ee DC 20013-7012, U.S.A. 34720D Durango, MEXICO 
peterson@si.edu; sorengr@si.edu yherrera@ipn.mx 
ABSTRACT 

Poa matri-occidentalis PM. Peterson & Soreng, sp nov, is described and illustrated. Poa matri-occidentalis subsp. 


matri-occidentalis is known from steep rocky slopes of Cerro Gordo, Durango, and Poa matri-occidentalis subsp. 
mohinorensis Soreng & P.M. Peterson, subsp. nov. is known from rocky cliffs of Cerro Mohinora, Chihuahua. The 


new species is morphologically similar to Poa kelloggii but differs by having leaf sheaths that are closed near the 
base 2/3 to 4/5 their length, entire ligules that are 3.5-6 mm long, nodding panicles, ovate lemmas with obtuse to 
acute apices, and non-visible rachilla internodes. A key distinguishing 11 species now reported from northern 
MAp 7 ] iq ] 


Is PlOVIGEC, 


RESUMEN 


s] Pay i bs 


Se Gescue e ilustra Poa matri- uceidentalis P.M. Peterson & Soreng, sp. nov. Poa matri 
-erro Gordo, Durango, y Poa matri- ‘cccldentalts 


subsp. mohinorensis Soreng & PM. Peterson, subsp. nov. se encuentra en la zona de laderas rocosos del Ce 

Mohinora, Chihuahua. La nueva especie es morfol6gicamente similar a Poa kelloggii pero difiere de ésta ce 
por tener las vainas de las hojas cerradas en la parte inferior de la base de 2/3 a 4/5 de su longitud, por tener 
igulas enteras de 3.5-6 mm de largo, paniculas recurvadas, lemas ovadas con apices agudos a obtusos y raquilla 
con entrenudos no-visibles. Se proporciona una clave para distinguir las 11 especies reportadas actualmente del 


— 


norte de México. 


Pod L. is the largest genus of grasses, including some 500-575 species that occur ina wide 
range of habitats from throughout the World (Soreng 1990; Gillespie & Soreng 2005). 
The genus is characterized by having rather small, multi-flowered spikelets, lemmas that 
are keeled, unawned, usually 5-nerved, commonly with web-like hairs emerging from 
the dorsal side of the callus, caryopses that are firm with lipid and a short hilum, lodi- 
cules that are broadly lanceolate with a lateral lobe, leaf sheaths closed above the base 
more than 1/20 the entire length, leaf blades that generally have two rows of bulliform 
cells (one on either side of the midnerve, these appearing as railroad tracks) and no addi- 
tional rows of bulliform cells, and blades commonly with naviculate (boat-shaped) api- 
ces (Soreng, in press). Poa has been divided into subgenera and sections and subsections 
for North America (Soreng 1998; Soreng in press), the New World (Soreng et al. 2003a), 
and the World (Gillespie et al. 2006; Zhu et al. 2006). 

Northern México (excluding Baja California and Baja California Sur) is not thought 
to be especially rich in Poa species. The following 11 species have been reported from the 
states of Aguascalientes (Ags), Chihuahua (Chih), Coahuila (Coah), Durango (Dur), Jalisco 
Jal), Nuevo Leon (NL), San Luis Potosi (SLP), Sonora (Son), Tamaulipas (Tamp), and 
Zacatecas (Zac): Pod annua L. (Ags, Chih, Coah, Dur, Jal, NL, SLP, Tamp, Zac), P. bigelovii 
Vasey & Scribn. (Chih, Coah, NL, Son), P. compressa L. (Coah), P. conglomerata Rupr. (= P. 
scaberula Hook. f.)[Coah is an error since a specimen (J.A. Garcia 49) cited in Beetle et al. 


SIDA 22(2): 905 — 914. 2006 


906 BRIT.ORG/SIDA 22(2) 


(1999) is = P. pratensis LJ, P fendleriana (Steud.) Vasey lincludes P fendleriana subsp. 
albescens (Hitche.) Soreng](Chih, Coah, Son, Zac), P infirma Kunth (listed as synonym of 
P.annua and probably with similar distribution), P mulleri Swallen (NL), P orizabensis 
Hitche. (NL possibly an error for CH. & M.T. Mueller 1248 from Cerro Potosi as deter- 
mined by A.A. Beetle =P. mulleri!, SLP-not yet verified by us from there), P. pratensis (Chih, 
Coah, NL), P. ruprechtii Peyr. includes P. sharpii Swallen as a synonym] (Coah, NL,), and 
P. strictiramea Hitche. (Chih, Coah, Dur, NL, Zac) [Beetle et al. 1999; Espejo Serna et al. 
2000; Hitchcock 1913]. Of the 10 species known from the region, P annua, P. compressa, P. 
infirma, and P. pratensis are introduced, and P. bigelovii, P fendleriana, P. mulleri, P. 
orizabensis, P ruprechtii, and P. strictiramea are native. The annuals, P annua and P 
infirma have been placed in Poa sect. Micrantherae Stapf, and P. bigelovii was placed in 
Poa sect. Homalopoa Dumort. (Soreng et al. 2003a). Of the remaining natives, P mulleri, P 
orizabensis, P. ruprechtii, and P. strictiramea have been placed in Poa sect. Homalopoa s.l, 
and P.fendleriana was placed in Poa sect. Madropoa Soreng (Soreng et al. 2003a). 

While making determinations of material in the field using Las Gramineas de 
Durango (Herrera Arrieta 2001) and then checking the herbarium at Centro 
Interdiciplinario de Investigacion para el Desarrollo Integral Regional (CIIDIR) in 
Durango, the first author recognized the unique features of a recent collection from Cerro 
Gordo. Upon further inspection of collections on loan from various herbaria for a com- 
plete revision of Poa in México, the second author discovered another, generally pubes- 
cent form, occurred near Cerro Mohinora, Chihuahua. The third author then contrib- 
uted the anatomical portion. We describe these two new forms as a new species of Poa 
with two subspecies. The new species with two subspecies is clearly aligned within sub- 
family Pooideae, tribe Poaeae, and subtribe Poinae (Soreng et al. 2003a, 2003b, 2005). 


Poa matri-occidentalis P.M. Peterson & Soreng, sp. nov. (Figs. la-c; 2¢e-1). Type: MEXICO. 
DURANGO: Sierra Madre Occidental, SW slope of Cerro Gordo just below twin rock outcrops (23°12'32.5"N- 
104°56'54.1"W), 3130-3200 m, 26 Sep 2005, P.M. Peterson 19145 & F Sdnchez Alvarado (HOLOTYPE: US! 
ISOTYPES CHDIR!, MEXUN). 


A Poa kelloggii Vasey vaginibus connatus 2/3-4/5 longe lateque, ligulis 3.5-6 mm longis integris, paniculus nutans, 
lematibus ovatis apicibus obrusus ad acutus, rachillibus no visibilis, recedit. 


Loosely caespitose and rhizomatous perennials with intra and extravaginal basal branch- 
ing. Culms 45-80 cm tall, solitary to several, erect or bases slightly decumbent, terete or 
weakly compressed; nodes 2-4, terete, glabrous and smooth, 1-3 exserted. Leaf sheaths 
mostly 3-14 cm long, closed about 2/3 to 4/5 their length, compressed, glabrous and 
smooth, or sometimes the lower ones retrorsely scabrous to puberulent and with ciliate 
collars, most sheaths 0.4-L.1 times as long as their blades; collars smooth or with a few 
hooks, glabrous or ciliate; ligules 3.5-6 mm long, membranous to hyaline, glabrous and 
smooth, or sometimes puberulent, apex obtuse to acute, entire; blades 2-6 mm wide, flat, 
abaxial nerves and margins lightly scabrous, adaxially smooth, glabrous throughout, 
apices narrowly prow-shaped; flag blades 12-22 cm long. Panicles 12-26 cm long, nod- 
ding, pyramidal, open, sparse with 24-85 spikelets, longest internodes 2.5-5.5 cm long; 
branches (1-)2(-3) at the lower nodes, longest 5.5-10 cm long bearing 3-15 spikelets, as- 
cending to spreading, lax, angled, the angles sparingly scaberulous. Spikelets 4-8 mm 
long, 1.8-2.7 mm wide, laterally compressed, greenish to stramineous, florets 2 or 3, per- 
fect, occasionally the upper floret is much reduced and sterile; lower rachilla internodes 
1-2 mm long, usually not visible, glabrous, smooth; glumes equal to subequal, lanceolate, 


PETERSON ET AL., POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 


= ee 
EF, = ga 
i 
o 
TL 
BAG Oye 


4 Le {Dd 


Al 


Fic. 1. Poa matri-occidentalis subsp. matri-occi (Peterson 19145 & Sd 
and blade. Poa tri identali } ie + rar AFJEOM-N 


lo). A. Habit. B. Inflorescence. C Sheath, ligule, 


Id). D. Sheath ligule and blade 


BRIT.ORG/SIDA 22(2) 


908 


Poa matri-occidentalis subsp. matri- 


glume. F. Upper glume. G. Lemma. H. Palea, dorsal 
K. lodicules. L. Young caryopsis 


Id). A. Spikelet. B. Floret 


AA-ND 
do). C. Spikelet. D. Floret. E. Lower 


ATCO 


Fic. 2. Poa matri 


Al 


hb 


occidentalis (Peterson 19145 & Sa 


at 


view. I, Palea, ventral view.J. Stamens, ovary 


7 


J 


7 


PETERSON ET AL., POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 909 


scabrous along the upper keel, distinctly keeled, 3-nerved, the lateral nerves often only 
evident near base, scaberulous along the nerves, apex acute to acuminate, of ten appear- 
ing mucronate because the margins are often involute, margins hyaline; lower glumes 3- 
5 mm long, lanceolate to ovate usually shorter than the upper; upper glumes 3.7-5.6 mm 
long, ovate to obovate, wider that the lower with more prominent nerves; calluses with 
cob-webby hairs, the hairs about 1/2 to 1/3 the length of the lemma; lowest lemmas 4.6- 
6.3 mm long, ovate, distinctly keeled, surfaces between nerves finely muriculate (with 
minutely pointed, whitish bumps on the short cells) or sometimes densely scabrous, 5- 
nerved, the lateral nerves moderately prominent, keel and lateral nerves scabrous, keel 
and marginal nerves glabrous or sometimes puberulent below, margins hyaline, apex 
obtuse to acute; lowest paleas 4.4-6 mm long, usually a little shorter than the lemmas, 
strongly laterally compressed with a deep adaxial furrow between the nerves, muriculate, 
scabrous along the nerves; anthers 2-2.2 mm long, yellow; lodicules 2, membranous; ovary 
glabrous. Caryopses 2.6-2.9 mm long, fusiform, light brownish. 

Phenology.—Flowering August through October. 

Comments.—The new species can be distinguished from Poa kelloggii by having leaf 
sheaths that are closed near the base 2/3 to 4/5 their length, entire ligules that are 3.5-6 
mm long, nodding panicles, ovate lemmas with obtuse to acute apices, and non-visible 
rachilla internodes. 


Poa matri-occidentalis subsp. matri-occidentalis (Figs. la—c; 2c-I; 3a-c; 4). 

Lower leaf sheaths glabrous and smooth; collars smooth or with a few hooks, glabrous: 
ligules glabrous and smooth. Lowest lemmas 4.7-5.4 mm long, surfaces between nerves 
finely muriculate (with minutely pointed, whitish-bumps on the short cells), keel and 
marginal nerves scabrous, glabrous throughout. 

Distribution and habitat—Known only from a single locality at Cerro Gordo, 
Durango, between 3130-3200 m where the species was found growing on steep, rocky, 
and grassy slopes beneath open forests of Pseudotsuga menziesii (Mirb.) Franco, Quercus 
sp., Pinus spp., with other associates, such as: Arctostaphylos pungens Kunth, Bromus 
carinatus Hook. & Arn., B. richardsonii Link, Aegopogon cenchroides Humb. & Bonpl. ex 
Willd., Trisetum viride (Kunth) Kunth, Festuca sp., Carex sp.,and Agrostis sp. 

Leaf anatomy.—Cross-sections were done by hand on re-hydrated material then 
mounted on temporary slides and stained with concentrated safranin. A diagram of a 
cross-section and abaxial leaf scrape were made with the aid of a camera lucida (Figs. 

aaa: 

Cross-section view (Fig. 3a-c.).—The lamina are V-shaped with primary, secondary, 
and tertiary vascular bundles of decreasing size. The primary vascular bundles are well 
differentiated into xylem with metaxylem, phloem, anda double bundle sheath (mestome 
and parenchyma bundle sheath). The ribs are rounded and furrows are <1/5 as deep as 
the width of the blade adaxially and abaxially. Abaxial projection of the midrib caused 
by inflated parenchyma cells and sclerenchyma is conspicuous. In half-the-width of the 
blade there is a single primary vascular bundle comprising the midvein; 3-4 additional 
primary vascular bundles; 3-4 secondary vascular bundles placed between consecutive 
tertiary vascular bundles; 7-8 tertiary vascular bundles placed between each consecu- 
tive primary and secondary vascular bundles; and two tertiary vascular bundles at the 
margin (i-iii-ii-iii-i-iii-ii-iii-i-iii-ii-iii-i-iii-ii-iii-iii). All vascular bundles are placed in the 
median layer of the blade and are round in outline. The xylem of the primary vascular 


910 BRIT.ORG/SIDA 22(2) 


n £hiad + £D, tr). id 


tpt oy - 1D. OVAL &. CAnrt Af, din\ 
7 


adaxial 


7 


Scale bar = 


‘ jie 
surface uppermost in all drawings. A. Primary vascular bundle. B. S Jary vascular bundle. C. T 
50um. 


Fic. 4. Abaxial leaf epidermis of Poa matri-occidentalis subsp. matri-occidentalis (Peterson 19145 & Sanchez Alvarado) showing long 
cells, silica bodies (dark), cork cells (clear), and th t ta. Scale bar = 


bundles contains two or occasionally three metaxylem vessels adjacent to the phloem, 
that are larger than the parenchyma bundle sheath cells, and sometimes one or two pro- 
toxylem vessels are located adaxially to the phloem. The mestome, or inner sheath, is 
always present in primary and secondary vascular bundles surrounding the xylem and 
phloem. The Mestome is composed of small cells with thick walls and interrupted 
abaxially by sclerenchyma fibers forming a girder in primary and secondary bundles, 
except in the central primary vascular bundle where it is continuous surrounding the 
bundle and the sclerenchyma is separated from the bundle by colorless cells and cen- 
tered at the very bottom of a conic abaxial base. The parenchyma bundle sheath, or outer 
layer, contains large, thin-walled cells; 10-12 cells are present on the continuous adaxial 
half circle of the primary vascular bundle sheath; 6-8 cells are present in the secondary 
vascular bundles; and the number of cells varies between three and four on each side of 
the interrupted tertiary vascular bundles. Chlorenchyma cells surround the bundles and 
are conspicuously continuous between the vascular bundles in a loose arrangement. Five 


PETERSON AND SORENG, POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 911 


to eight sclerenchyma fibers are present adaxially in all bundles. Five to eight rows of 
sclerenchyma fibers form the abaxial girders of the primary and secondary vascular 
bundles, which widen near the epidermis and narrow toward the vascular bundle. Deep 
fan-shaped rows of bulliform cell are located adaxially near the central part of the blade 
between the central primary vascular bundle and two lateral tertiary vascular bundles: 
these are continuous to adaxial epidermal cell walls. 

Epidermis in abaxial view (Fig. 4).-Costal and intercostals zones are clearly distin- 
guishable. Three to six rows of short cells are found in the costal zone, each separated by 
a row of long cells. Irregularly-shaped silica bodies are alternate and adjacent to short 
cells of the costal zone. Cork cells are less frequent and the same shape as the silica bod- 
ies. Fifteen to seventeen rows of long cells are common in the intercostal zone. Long cells 
are 8-10 times longer than wide with wavy cell walls. No papillae or hairs are present. 
Low dome-shaped stomata are infrequent and found in single row. Parallel subsidiary 
cells are rounded in outline. A few antrorse bristles are present at foliar margin. 


Poa matri- cece subsp. mohinorensis Soreng & PM. Peterson, subsp. nov. (Figs. 1d; 
2a, ).T EXICO. CHIHUAHUA: Municipio de Guadalupe y Calvo, Sierra Nees ee east Nside of 
Cerro <rnaah ca. 13 mi SW of Guadalupe y Calvo, open pine-fir woods with scattered spruce, noe 
vertical, N-facing rock wall, very moist with me bryophytes and rich herbaceous flora (25°57' 
107°03'W), 2950 m, 20 Aug 1988, G. Nesom 6475 & A. McDonald (HOLOTYPE: TEX!: ISOTYPE ARIZ). 


A Poa matri-occidentalis subsp. matri-occidentalis vaginae disatalibus et lemmatae carina puberulens, 
; an fee or 
terner 1 abris, differt 


Lower leaf sheaths retrorsely scabrous to puberulent; collars ciliate; ligules puberulent. 
Lowest lemmas 4.6-6.3 mm long, surfaces between the nerves densely scabrous, keel and 
marginal nerves scabrous and puberulent below. 

Distribution and habitat.—Known only from Cerro Mohinora, Chihuahua between 
2950-3130 m, on moist, rocky cliffs and ledges associated with open forests of Pinus, Abies, 
and Picea. 


Additional specimen examined. MEXICO. Chihuahu lupe y Calvo, Sierra Madre Occiden- 
~ on high ledges on summit of Cerro Mohinora OStS7N- 107°03 W), 10, 000- 10,300 ft, 16-17 Oct 1959, D.S. Correll 
177 & H.S. Gentry (LL). 


DISCUSSION 


Based on morphological similarity to Poa kelloggii, endemic to California, P. matri- 
occidentalis is tentatively placed in Poa sect. Sylvestres V.L. Marsh ex Soreng. Poa sect. 
Sylvestres as treated for the genus in the Flora of North America, includes seven species, 
all of which are endemic to forests in the U.S.A. and southern Canada (Soreng in press). 
[Poa sect. Sylvestres slightly modified from Soreng in press] “Plants perennial; usu- 
ally non-rhizomatous and non-stoloniferous, sometimes shortly rhizomatous, usually 
loosely tufted, infrequently densely tufted. Basal branching mainly pseudointravaginal, 
or sometimes mainly extravaginal. Culms 20-126 cm, terete or weakly compressed. 
Sheaths closed (1/20)1/3 to about their full length, terete or weakly keeled, basal sheaths 
readily deteriorating; ligules 0.1-3(-4) mm long, smooth or sparsely scabrous, truncate 
to obtuse, entire or lacerate, smooth or ciliolate; blades smooth or scabrous, glabrous, api- 
ces narrowly prow-shaped. Panicles 4-36 cm long, erect or lax, pyramidal or lanceoloid, 
usually sparse, lower rachis internodes usually longer than (2-)3 cm, nodes with 1-10 
branches; branches ascending to spreading or eventually reflexed, lax or straight, angled, 
angles scabrous, spikelets confined to distal 1/5-1/3(-1/2). Spikelets 2.5-8.2 mm long, 


912 BRIT.ORG/SIDA 22(2) 


laterally compressed: florets (1-)2-5(-6), normal; rachilla internodes smooth, usually gla- 
brous, sometimes hairy, hairs to 0.2 mm long. Glumes distinctly keeled, scabrous on the 
keel and nerves; calluses terete or slightly laterally compressed, usually dorsally webbed, 
sometimes glabrous; lemmas 2.1-5 mm long, lanceolate to broadly lanceolate, distinctly 
keeled, glabrous or hairy, apices with narrow, clear or white margins; palea keels sca- 
brous, glabrous or with hairs over the keels; anthers 3, 0.4-2(-2.6) mm long.” 
Based on DNA sequence analyses, Poa sect. Sylvestres is the earliest diverging section 
in the genus, if Arctopoa (Griseb.) Prob. is excluded, as we now believe it should be 
(Gillespie et al. in prep.). However, the new species differs in some respects from all or 
most of the other species in the section. Like P. kelloggii, P matri-occidentalis is short 
rhizomatous and its basal branching is primarily extravaginal. Poa matri-occidentalis 
differs from all the other species in the section by having longer ligules (3.5-6 mm long). 
If the new species is confirmed to belong to this section (viz. DNA sequence analysis) it 
would be the first report from outside the U.S.A. and Canada. Our only hesitation to plac- 
ing the new species in this section is that there is some morphological overlap with Poa 
sect. Homalopoa sl. and with the P. nervosa (Steud.) Vasey complex of Poa sect. Mad ropoa, 
and currently, there are no DNA data for the new species. In Poa sect. Madropoa, through 
subsp. mohinorensis, P matri-occidentalis most closely approaches P. tracyi Vasey from 
New Mexico and Colorado U.S.A. (as noted on the 1989 annotation by J.R.and C.G. Reeder 
on Nesom 6475), and “P. ruprechtii Peyr.” (as that name is applied to material from the 
Sierra Madre Oriental, México). Poa matri-occidentalis differs from both these species by 
having muriculate or short-pubescence on the lemmas and only perfect flowers. The new 
species differs from “P. ruprechtii” by having longer glumes that are also proportionally 
longer relative to the adjacent lemmas and by fewer florets per spikelet. We suspect the 
name P ruprechtii Peyr. s. str. may be misapplied in northern México’s Sierra Madre Ori- 
ental as material called this by Hitchcock (1913) and Fournier (1886), from the region of 
the type locality, in the State of México and Distrito Federal, differs in having short an- 
thers,and more indurate and prominently nerved lemmas with relatively longer glumes, 
but we have yet to locate and examine a type specimen of P ruprechtii. If our suspicion is 
correct, the Sierra Madre Oriental plants probably represent an additional new species. 
ey to all known or reported species from northern México follows. In this key 
all measurements were taken from the lowest, one or two florets in the best-developed 
spikelets. 


KEY TO THE SPECIES OF POA IN NORTHERN MEXICO 


1. Plants annual; palea keels pubescent in part. 
2. Callus with cobwebby hairs; panicles contracted P. bigelovii 
2. Callus glabrous; panicles open. 
3. Anthers 0.6—1 mm long 
3. Anthers 0.2-0.5 mm long 
1. Plants perennial; palea keels scabrous only. 
4. Anthers 0.5—1 mm long callus webbed. 
5. Lemmas glabrous or very sparsely puberulent on the keel base; spikelets broadly ovate 
P. orizabensis 
5. Lemmas short villous on the keel and lateral nerves and sometimes puberulent between 


P,annua 
P.infirma 


the nerves; spikelets broadly lanceolate (of Distrito Federal and State of México) P. ruprechtii 
4. Anthers 1.2-3 mm long; callus glabrous or pubescent. 
6. Plants without rhizomes or lateral tending shoots, densely tufted, vegetative shoots mainly 


intravaginal. 


PETERSON AND SORENG, POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 


7. Upper glumes (3-)5-7-nerved; lemmas pubescent on lower half of the keel; longest 
branches up to 3.5-3.8 cm long; plants 20-42 cm ta P. mulleri 

7. Upper glumes 3-nerved; lemmas glabrous or very sparsely puberulent on the keel and 

Ha nerves near the base; longest branches up to 10 cm long; plants mostly (20-) 
40-10 tall P. strictiramea 

6. Plants fe ene or lateral tending extravaginal shoots, densely to loosely tufted, veg- 

etative shoots intravaginal “ve extravaginal or mostly Oavag 
8. Panicles contracted, fairl f bl 


d, leaf blades | lig! ily reduced O! absent; Ppopu- 
lations dioecious or pistillate; callus glabrous. 


9. Lemmas glabrous P.fendleriana ssp. albescens 
9. Lemmas BUDE cent on une ie and marginal nerves P. fendleriana ssp. fendleriana 
. Panicles loosely contract leaf blades well-developed; plants 
open; callus, at least of proximal 


all perfect flowered, or, if piiliane in part then panicl 


lemmas, with co eee hairs or giebie ous 
10. Lemmas aa 


sparingly puberulent near the base of the keel; upper glumes 
3.7-5.6mm ae ligules obtuse to acute, 3-6 mm lon 

Sheaths of lower leaves retrorsely scabrous to puberulent; collars ciliate; lem- 
mas densely scabrous between the nerves, keels and marginal nerves 
puberulent below 


Sheaths of lower leaves smooth, gla coll 


= 


Vi 
matri-occidentalis sso. mohinorensis 
rs smooth or with a few hooks, 
glabrous; lemmas finely lin pee ae us nerves, keel and marginal 
nerves glabrous below 


— 
— 


ri-occidentalis ssp 


oS 


P.m matri-occidentalis 
Lemmas distinctly pubescent on the keel and a nerves; ligules Glnese O 
obtuse, 1-3 mm long. 


12. Culms and nodes strongly compressed, keeled 
12. Culms and nodes terete, or culms weakly compressed 
13, peal inteiedes nosy nleaen ree view; piel Mlesuy os . mm 


P. compressa 


pie entire, usually 1-2 mm ce P. pratensis 
Rac 


ee. 


A 5-8 | pee £ ( 


least the bona most ones) with hairs between the keel and ete 


nerves near the base; |i obtuse, irregularly dentate, to 3 mm long (0 
Sierra Madre Oriental) 


“P.ruprechtii” 
ACKNOWLEDGMENTS 


We wish to thank the Smithsonian Institutions Restricted Endowment Fund and the 
Biodiversity, Surveys, and Inventories Program at the National Museum of Natural His- 
tory for supporting the fieldwork. Appreciation is extended to Alice R. Tangerini for pro- 
viding the illustrations; Patricia GOmez Bustamante for correcting the Spanish resumen 


Dan Nicolson for correcting the Latin diagnoses; and Jesus Valdés-Reyna and Adolfo 
Espejo-Serna for reviewing the manuscript. 


REFERENCES 
Beette, A.A., G. ViLLecas Duran, A. Botanos Mepina, J.A. MIRANDA SANCHEZ, L. ARAGON MetcHor, M.A. Vergara 
Batalla, A. CHiMAL HerNANoez, M.M. Castitto BabiLto, O.M. GALAN Gar A, J.L. VILLALPANDO Prieto, M. LIZAMA 
Manriaue, J. VALDES-Reyna, E. MANRIQUE DE SKENDZIc, and A.M. Ropricuez Ropricuez. 1999. Las Gramineas 
de México, Tomo V. Secretaria de ieee ltt ae y Desarrollo Rural, Distrito Federal, México 
Espeso Serna, A., A.R. Lorez-Ferrari, and J. Vacoés-Reyna. 2000. Poaceae. In: A. Espejo-Serna & A.R. L6pez- 
Ferrari,eds.Las Monocotiledéneas Mexicanas:una Sinopsis Floristica, Partes IX—X|. Consejo Nacio- 
nal de la Flora de México, A.C., Universidad Aut6noma Metropolitana-|ztapalapa, and Comision 
Nacional para el Conocimiento y uso de la Biodiversidad, México, D.F. Pp. 108-236 
Fouenier. E. 1886. Mexicanas plantas. Pars secunda: Gramineae 2:1-160. 


914 BRIT.ORG/SIDA 22(2) 


Gitespie, LJ, A. ARCHAMBAULT, aNd RJ. SoreNc. 2006. Phylogeny of Poa (Poaceae) based on trnT-trnF 
sequence data: major clades and basal relationships.. In: J.T. Columbus, E.A. Friar, J.M. Porter, L.M. 
Prince,and M.G. Simpson, eds. Monocots: comparative biology and evolution, 2 vols.Rancho Santa 
Ana Botanic Garden, Claremont, California. Pp. xx—xx. 

Giiespic, L.J.and R.J.Sorena.2005.A phylogenetic analysis of the bluegrass genus Poa based on cpDNA 
restriction site data. Syst. Bot. 30:84-105. 

Herrera Arrieta, Y.2001.Las Gramineas de Durango. Instituto Politécnico Nacional and Comision Na- 
cional para el Conocimiento y Uso de la Biodiversidad, Durango, México. 

HitcHcock, A.S. 1913. Mexican grasses in the United States National Herbarium. Contr. U.S. Natl. Herb. 
17(3):181-389. 

Sorene, R.J. 1990.Chloroplast-DNA phylogenetics and biogeography in a reticulating group: study in 
Poa (Poaceae). Amer. J. Bot. 77:1383-1400. 

Sorene, R.J.1998.An infrageneric classification for Poa in North America, and other notes on sections, 
species, and subspecies of Poa, Puccinellia, and Dissanthelium (Poaceae). Novon 8:187-202. 

Sorena, RJ. In press. Poa L. In: Barkworth, M.E., KM. Capels, S. Long, M.B. Piep, eds. Magnoliophyta: 
Commelinidae (in part): Poaceae, part 1.Flora of North America North of Mexico, volume 24. Ox- 
ford University Press, New York. Pp. xx-xx. 

Sorene, R.J.G. Daviose, BM. Peterson, F.O. ZuLoaca, E.J. Jupziewicz, T.S. Fitcueiras, and O. Morrone. 2005. Cata- 
logue of New World grasses (Poaceae). http://mobot.mobot.orgW3T/Search/nwgc.htmland Clas- 
sification of New World Grasses. http://mobot.mobot.org/W3T/Search/nwaclass.html. 

Soreno, R.J., L.Giussani, and M. Necrito. 2003a. Poa. |n: Soreng, R.J.,P.M. Peterson, G. Davidse, E.J. Judziewicz 
FO. Zuloaga, TS. Filgueiras, and O. Morrone, eds. Catalogue of New World grasses (Poaceae): IV. 
Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:505—580. 

Soren, R.J., BM. Peterson, G. Daviose, EJ. Juoziewicz, F.O. ZULOAGA, T. S. FitGueiras, and O. Morrone. 2003b. 
Catalogue of New World grasses (Poaceae): IV: subfamily Pooideae. Contr. U.S. Natl. Herb. 48: 


Zuu,G,, L. Liu, RJ. Sorenc, and M.V. Otonova. 2006. Poa.In: Wu, Z.Y,,P.H.Raven, and D.Y. Hong, eds. Flora of 
China: Poaceae (Gramineae), volume 22. Science Press, Beijing & Missouri Botanical Garden Press, 
St. Louis. Pp. 257-309. 


BROMUS AYACUCHENSIS (POACEAE: POOIDEAE: BROMEAB), 
A NEW SPECIES FROM PERU, WITH 
A KEY TO BROMUS IN PERU 


Jeffery M. Saarela Paul M. Peterson 
Department of Botany, and Department of Botany 
UBC Botanical Garden and Centre for Plant Research National Museum of Natural History 
Sek ee Columbia Smithsonian ee 
a C., CANADA Washington, DC 20013-7012, U.S.A. 
ee er tice peterson@si.edu 


Nancy F. pelos ee 
Rancho Santa An arde 
1500 North Co ig Avenue 
Claremont, California, 91711-3157, U.S.A. 
ee 


ABSTRACT 


Bromus ayacuchensis, a new species from Departamento Ayacucho, Peru, is described and illustrated. The new 
species is similar to species in Bromus sect. Ceratochloa, particularly B. ce ae because of its strongly keeled 
ase, adaxially pilose ligules 3-6.6 


] 


lemmas. It differs from : cebadilla by having densely pil the 

mm long, lower glumes I-veined, upper glumes 3-veined, pilose lemmas es y along the margins and near 
the apex), lemma margins that are tightly involute on lower 1/5-1/4, and lemma awns 3-5.5 mm long. Phyloge- 
netic analyses of sequence data from the internal transcribed spacer regions of nuclear ribosomal DNA place B. 
ayacuchensis in a clade with species of sect. Ceratochloa, whereas analyses of plastid data from the trnL intron 
and the 3-end of ndhF indicate that B. ayacuchensis is closely related to this clade. Confirmed records are pro- 
vided for B. modestus and B. cebadilla in Peru, and B. coloratus and B. flexuosus are reported for the first time in 
Peru. A key to the 12 species of Bromus now known from Peru is included. 


RESUMEN 


Se describe y se il hensis Saarela & P.M. Peterson, sp. nov, una nueva especie del Departamento 


acucho, Pert. La nueva ponacic es similar a las especies de Bromus secc. Ceratochloa, pa tnculnentc B. 


cae | 


Cea porque tiene lemas muy carenadas. Se diferencia de B. cebadilla por tenet 
cerca de la base, ene 3-6.6 mm de largo y pilosas en la parte ventral superior, glumas inferiores con if nervio, 
glumas sup 3 nervios, lemas pilosos (especialmente a lo largo de los margenes y cerca del apice), los 
margenes del lema son firmemente involutos en la porcioén 1/5-1/4 inferior del lema y aristas de 3-5.5 mm de 
largo. Los analisis filogenéticos de los datos de ae secuencia de ADN de las Repiories transcritas internas del 
espaciador ribosomico nuclear colocan a B. ayact n clado con las especies de la sect. Ceratochloa, 
mientras que los analisis de los datos del intrén del ee del trnL y de los extremos : del ndhF indican que B. 
ayacuchensis esta intimamente pacionade pero no es parte de este clado. Proporcionamos ejemplares de 
especimenes B. modestus y B.cebadilla pro lel Pert, y B.coloratus y os XUOSUS ae. por primera 
vez en Peru. Se incluye la clave para las 12 especies de Bromus conocidas en Per 


Bromus L. (Poaceae: Pooideae: Bromeae) is a large genus of grasses with over 160 species 
distributed throughout temperate regions of the world (Clayton & Renvoize 1986). Some 
species are important forage grasses (e.g., B. inermis Leyss.; B. auleticus Trin. ex Nees), and 
many are highly aggressive, invasive weeds (e.g., B. diandrus Roth; B. madritensis L; B. 
tectorum L.; Pavlick 1995 

Bromus is distinguished from other grass genera by the combination of leaf sheath 


SIDA 22(2): 915 - 926. 2006 


916 BRIT.ORG/SIDA 22(2) 


margins that are connate (closed) for most of their length, awns that are inserted subapi- 
cally on the lemmas (except in B. lepidus Holmb. and B. bidentatus Holmstr6m & H. 
Scholz; Holmstrom & Scholz 2000), hairy apical bilabiate-appendages of the ovary, and 
simple starch grains (Wagnon 1952; Smith 1970; Clayton & Renvoize 1986). 

Bromus is diverse morphologically and has been divided variously into segregate 
genera[Anisantha C. Koch, Boissiera Hochst. ex Steud., Bromus, Bromopsis (Dumort.) Fourr, 
Ceratochloa P. Beauv, Nevskiella Kreczetowicz & Vvedensky, and Trisetobromus Nevskil, 
subgenera[Bromus, Stenobromus (Hack.), Festucaria Gren. & Godr., Ce ratochloa (P. Beauv.) 
Hack., Nevskiella (Kreczetowicz & Vvedensky) Kreczetowicz & Vvedensky, and 
Neobromus Shear: Stebbins 1981] or sections [Bromus, Triniusia (Steud.) Nevski., Boissiera 
(Hochst. ex Steud.) P.M. Sm., Genea Dumort., Bromopsis Dumort., Ceratochloa (P. Beauv.) 
Griseb., Nevskiella (Kreczetowicz & Vvedensky) Tournay, and Neobromus (Shear) Hitche.; 
Smith 1970]. Most recent workers use a sectional classification (e.g., Clayton & Renvoize 
1986: Pavlick 1995; Planchuelo & Peterson 2000; Pavlick et al. 2003). 

Molecular studies (Pillay & Hilu 1990, 1995; Ainouche & Bayer 1997; Saarela et al. in 
press) support the monophyly of some of these sections (e.g, Bromus sects. Bromus, Genead, 
Ceratochloa), and have indicated that others comprise several distinct lineages (e.g., 
Bromus sect. Bromopsis). Incongruence between plastid and nuclear ribosomal gene trees 
has provided tentative insight into past hybridization events in the genus (Saarela et al. 
in press). The available classification schemes (e.g., Smith 1970) do not satisfactorily reflect 
evolutionary relationships in the genus as understood currently (Saarela et al. in press). 

Approximately 80 native and introduced species of Bromus are recognized currently 
in North, South, and Central America (Pavlick et al. 2003). Many regional treatments of 
Bromus have been published (Wagnon 1952; Soderstrom & Beaman 1968; Camara 
Hernandez 1978; Pinto-Escobar 1981, 1986; Matthei 1986; Pavlick 1995; Gutiérrez & Pensiero 
1998; Planchuelo & Peterson 2000; Pavlick et al. 2003), cytological, genetical, and ana- 
tomical studies are revealing new information (e.g., Martinello & Schifino- Wittman 2003; 
Ramos et al. 2002; Fuet al. 2005; Tuna et al. 2005, 2006), taxonomic studies are clarifying 
species circumscriptions (e.g., Naranjo et al. 1990; Planchuelo 1991; Naranjo 1992; Peter- 
son et al. 2002; Oja et al. 2003; Massa et al. 2001, 2004; Saarela et al. 2005) and new taxa 
are being described (e.g., Matthei 1986; Renvoize 1994; Peterson & Planchuelo 1998). 

Ona recent trip to Peru, the second and third authors collected several unusual indi- 
viduals of Bromus with strongly keeled (laterally flattened) lemmas, l-veined lower 
glumes, and 3-veined upper glumes. Strongly keeled lemmas are characteristic of sect. 
Ceratochloa, but all known species from this section have 3-9-veined lower glumes and 
5-9-veined upper glumes (Smith 1970). Known species of Bromus that have 1- and 3-veined 
lower and upper glumes, respectively, have lemmas that are wide and short, rounded, or 
slightly keeled (Smith 1970). A review of the taxonomic literature and specimens in the 
US National Herbarium has indicated that no individuals of Bromus with this combina- 
tion of characters have been described previously. We name and describe these recent 
collections as a new species, Bromus ayacuchensis. We discuss the phylogenetic place- 
ment of the new species in the genus based on its morphology and DNA sequence data 
from the internal transcribed spacer (ITS) regions of nuclear ribosomal DNA, the plastid 
trnL(UAA) intron, and the 3’-end of the plastid ndhF gene. 


MATERIALS AND METHODS 


The morphological description of the new species is based on material from three collec- 


SAARELA ET AL., A NEW SPECIES OF BROMUS FROM PERU 917 


tions (see below). Fifteen specimens were measured for the morphological description. 
To characterize leaf anatomy, leaf blades were field collected, and segments of about 5 
mm in length from the middle of the second leaf blade below the inflorescence of the 
flowering culm were fixed in FPA (formalin: propionic acid: ethyl alcohol: water; 2:1:10:7). 
Leaf blades were desilicified in 30%-hydrofluoric acid for 48 hours. Leaf blade segments 
were dehydrated, embedded, sectioned, and stained following the methods described in 
Columbus (1999). Terminology used in the anatomical description follows Metcalfe (1960) 
and Acedo & Llamas (1999). 

DNA was obtained from field collected leaf material of two collections (Peterson 
16452 & Refulio-Rodriguez and Peterson 18212 & Refulio-Rodriguez). Voucher specimens 
are deposited in US. Sequence data were obtained for B. ayacuchensis from the plastid 
trnL(UAA) intron, the 3’-end of the plastid ndhF gene, and the internal transcribed spacer 
(ITS) regions of nuclear ribosomal DNA, following the protocol in Saarela et al. (in press). 
Sequences for all regions from both individuals were identical, thus a single consensus 
sequence was generated for each region. Sequence data have been deposited in GenBank 
with the following accessions: (ITS: DQ676867; 3’ ndhF: DQ676868; trnL intron: 
DQ676866). The new sequence data were added to an existing matrix that includes se- 
quence data from 46 species of Bromus that represent a large proportion of the morpho- 
logical and geographical diversity in the genus (Saarela et al. in press). Heuristic searches 
using parsimony conducted independently on the nuclear ribosomal DNA and com- 
bined plastid data sets with 100 random starting trees, tree-bisection-reconnection (TBR) 
branch swapping, and all character and character-state changes equally weighted. Branch 
support was assessed using maximum parsimony bootstrap analysis (Felsenstein 1985) 
from 500 replicates using the heuristic search option, with one random starting tree, TBR 
branch swapping, and MaxTrees set to 500 per replicate. 


RESULTS AND DISCUSSION 


Morphological and molecular data suggest that B. ayacuchensis is similar to species clas- 
sified in Bromus sect. Ceratochloa. Bt ayacucl I the strongly laterally com- 
pressed spikelets with species in sect. Ceratochloa, and the nuclear ribosomal trees in- 
clude it in a clade with species of Bromus sect. Ceratochloa (BS=92%; Fig. la). Bromus sect. 
Ceratochloa is native to North and South America, characterized by a perennial habit, 
ovate-lanceolate strongly laterally compressed spikelets, 3-5-veined lower glumes, 5-7- 
veined upper glumes, and strongly keeled lemmas (Smith 1970). All taxa in the section 
are polyploids (octo-, hexa-, and 12-ploid) (Stebbins 1981; Pavlick 1995). In the first com- 
plete synopsis of Bromus in South America, Planchuelo & Peterson (2000) recognized 
nine species in sect. Ceratochloa. A recent taxonomic study has proposed the lumping of 
several hexaploid taxa from Patagonia [B. catharticus Vahl, B. cebadilla Steud. (syn. = B 
stamineus E. Desv.), B. coloratus Steud., B. lithobius Trin., B. mango E. Desv., and B. tunicatus 
Phil] into one morphologically variable species, B. catharticus (Massa et al. 2001, 2004). 
Other species of sect. Ceratochloa recognized currently in South America include B. 
bonariensis Parodi & J.A. Camara, an endemic of Provincia Buenos Aires, Argentina, and 
B. striatus Hitchc., an endemic of Peru (Planchuelo & Peterson 2000; Pavlick et al. 2003). 
These species were outside of the geographic scope of the studies by Massa et al. (2001, 
2004), but both have been considered conspecific with B. catharticus in the past (Pinto- 
Escobar 1986; Planchuelo 1991). Among these taxa, B. dyacuchensis is most similar to B. 
cebadilla, a species known from Argentina, Chile, Peru, and the USA (Pavlick et al. 2003). 


B. anomalus 


Bui 

B le an ail § 
B. nottowayanus 
B, pubescens 
B. attenuatus 

B. grandis 

B. exattatus 


Bromopsis 


B. scoparius 
B, japonicus 


BRIT.ORG/SIDA 22(2) 


B. anomalus 
B. attenuatus 
B. ciliatu 
B. pubescens 
B. psendolaevipes 
B. pumpellianus 
B. ramosus 


B. erectus 
B, riparius 
B, exaltatus 

B. frondosus 


B, laevipe Bromopsis 


B. pectinatus 1 Bromus ; 
B. danthoniae B. grant is 
5 ™ B. pseudodanthoniae B. mucroglumis 
B. texensis es Inernus 
8. frondosus t 
B. suksdorfii B. lanatipes 
natipes 4 B. richardson 
arteri Bromopsis 4 B. texensis 
B. pseudolaevipes B. korotkovi 
B. mucroglumis igdumis 
B. richardsonii f 
B TGUNOSUS ; B. suksdorfii 
. brachyanthera | Bromopsis gh (7 2 seeparins 
. marginatus . B. japonicus 
B. coe anus B. pseudodanthoniae Bromus 
ams 8% B. danthoniae 
—— B. subvewtinus Ceratochloa B. diandrus | Genea 
B. catharticn Dinas 
a -pectinats — | Bromus 
B. cebadilla A naaidhileisis-sub hens) 
B cocina 5 oi inddrifensis SUDSP. rubeNs 
fs R / I rn 3 1 
B. ayacuchensis . Genea 
= B. trinil madritensis subsp. rubens 2 
0 B. gunckeili | Neobromus 
70 B diandrus . 
100 B, madritensis subsp rubens | _ Bromopsis 
B. madritensis subsp. rubens Genea 
B. madritensis subsp. rubens 
B. erectus Bromopsis 
B. riparius 
B. inermis oN 
75 B. pumpellianus 
Be daratkoyt Ceratochloa 
rm 8. ances) Bromopsis 
100 B. pflanzii 
B. modesta 
68 B, pellinus 
100 —. B. densus 
8 dolich oe Neobromus 
Triticum aestivum 
m Horde um i alec ed Hordeum vulgare 
estuca brevighumi Festuca breviglum 
== 5 changes — 10 changes 
1 fp L L 4 PP £, (A\ eb | by 4 (IT 
egi ons of 


Fic. iti 
eee ribosomal DNA. Ine position of ‘ arene! in oa) tree is indicated with an open arrow. Bootstrap support alte are 
Tradition- 


ally recognized sections in Bromus (sensu Smith 1970) 


and 3’-end of ndhF 


Bromus ayacuchensis differs from B. cebadilla by having densely pilose culms near the 
base, adaxially pilose ligules 3-6.6 mm long, lower glumes l-veined, upper glumes 3- 
veined, pilose lemmas (especially along the margins and near the apex), lemma margins 
that are tightly involute on lower 1/5-1/4, and lemma awns 3-5.5 mm long. The leaf blade 
of B. ayacuchensis (Fig. +) is anatomically similar to the leaf blade of B. catharticus (the 
only taxon in sect. Ceratochloa tor which anatomical data exist; Acedo & Llamas 1999). It 
has a well-developed keel, Wee: ecemnaee bulliform cells, and lower-order vascular 


bundles associated with s ma 
Morphological and molecular data also identify some eierenecs between B. 
: 12 LE 


ayacuchensis and species classified in sect. Ceratochloa. Bromus ayacuch from 
taxa classified in sect. Ceratochloa by its 1- and 3- veined lower glumes. In the plastid 
trees, B. ayacuchensis is weakly supported as the sister group of a clade that comprises 
Bromus sect. Ceratochloa and B. brachyanthera, and these taxa are part of a larger clade 
that also includes species of sect. Neobromus (Fig. lb). Among South American Bromus 


SAARELA ET AL., A NEW SPECIES OF BROMUS FROM PERU 919 


species, l- and 3-veined lower and upper glumes are found in taxa traditionally classified 
within Bromus sections Bromopsis, Genea, and Neobromus (Smith 1970). The two species 
classified in ie closely-related sect. Neobromus are distinguished from B. ayacuchensis 
by their narrowly elliptic spikelets, lemmas with a deep apical sinus, and geniculate awns 
(Smith 1970; Matthei 1986). Species of sect. Genea in South America, which are all intro- 
duced, are easily separated from B. ayacuchensis by their oblong or wedge-shaped spike- 
lets and narrow and elongate lemmas with long awns (Smith 1970). Bromus sect. Bromopsis 
is likely a paraphyletic assemblage of species (Saarela et al. in press) that have 1(-3)- and 
3(-5)-veined lower and upper glumes, similar to B. ayacuchensis. The strongly laterally 
compressed spikelets and lemmas of B. ayacuchensis separate it from this artificial group 
of species, which have spikelets and lemmas that are terete and dorsally flattened or some- 
times slightly keeled (Smith 1970), although the plastid and nuclear molecular data indi- 
cate that one species of sect. Bromopsis from South America, B. brachyanthera, is closely 
related to B. ayacuchensis and sect. Ceratochloa (Figs. la, b). The molecular data and mixed 
morphological characters suggest that B.ayacuchensis might have originated via hybrid- 
ization between a species of sect. Ceratochloa and a species of the non-monophyletic sect. 
Bromopsis. Sampling of additional South American species of sect. Bromopsis will be nec- 
essary to determine if any are part of the clade that includes sects. Neobromus, Ceratochloa, 
B. brachyanthera and B. ayacuchensis 

Nine species of Bromus (B. berteroanus Colla; B. catharticus; B. cebadilla; B. lanatus 
Kunth; B. modestus Renvoize; B. pitensis Kunth; B. segetum Kunth; B. striatus Hitchc.; B. 
villosissimus Hitchc.) are currently known and verified by us from Peru (Planchuelo & 
Peterson 2000; Tovar 1993). Bromus carinatus Hook. & Arn. and B. inermis Leyss. have 
also been listed for Peru but these records are doubtful (Brako & Zarucchi 1994). Bromus 
cebadilla has been reported from the country previously without citation of specimens 
(Renvoize 1998; Pavlick et al. 2003). We provide confirmed records of this taxon in Ap- 
pendix 1. Bromus modestus is a recently described species (Renvoize 1994), and we report 
additional collections from Peru in Appendix 1. We have also collected B. coloratus and B. 
flexuosus Planchuelo in Peru (Planchuelo 1983), and report these collections here for the 
first time (Appendix 1). With the discovery of B. ayacuchensis, and confirmed records of 
B. coloratus and B. flexuosus, the number of Bromus species known from Peru is 12. A new 
key to Bromus in Peru is provided below. 


~— 


TAXONOMIC TREATMENT 


Bromus items Saarela & PM. Peterson, sp. nov. (Fig. 2). Type: PERU. DeParTAMENTO 
AYACUCHO: Provincia Lucanas, 12 km E of Puquio on road towards Cuzco, (14°4119.2" S Lat, 74°04'28.3" W 
Long), coe , among large boulders with Berberis (Berberidaceae), above meadow with small creek, 11 
Mar 2002, PM. nee 16452 & NE Refulio-Rodriguez (HOLOTYPE: US! ISOTYPES: K!, MO!, UBC!, USM). 


A Bromo cebadilla Steud. culmis Sah basilaribus, adaxialibus pilosis 3-6.6 mm longis ligulis, inferioribus 1- 
nervatis glumis, superioribus 3-nervatis glumis, pilosis lemmatibus (praecipue apud margines et apicem 
lemmatum marginibus inferioribus je 1/4 involventibus, lemmatum aristis 3-5.5 mm longis, recedit. 
Loosely caespitose perennials, rhizomes short with extravaginal shoot initiation. Culms 
(30-)50-120 cm tall, erect to slightly decumbent near base, pilose, densely pilose near 
base; internodes somewhat glabrous to sparsely pilose, retrorsely pilose just above dark 
brownish nodes. Sheaths 8-20 cm long, mostly longer than internodes below and shorter 
than internodes above, closed on lower 2/3-3/4, retrorsely pilose, hairs 0.7-1.6 mm long; 
margins smooth; cataphylls and lower sheaths densely pilose, often shredding. Ligules 


BRIT.ORG/SIDA 22(2) 


Nae ’ 
= } Be — 
—_ 
S Se 
y a 
ie in 


fe] 
en \ _ 
p \. 


N 


ae — 


SS. 
\ A RTAMCER I] 2004 


} N\ 
We AOS, 
ANN 


Fig. 2. Bromus ayacuchensis (Peterson 16452 & Refulio-Rodriquez). A. Habit. B. Inflorescence. C. Sheath, ligule, and base of blade. D. 
Spikelet.E. Lower glume, dorsal view. F. Upper glume, dorsal view. G. Lemma, dorsal view. H. Floret, ventral view. |. Floret, lateral view. 
J Q £41 ey | kill - K p.l es a Vas T J ventral view I Dp. *aL hin J 


J 


dorsal view. M. Lodicules, stamens, and pistil. N. Lodicules. 0. Gynoecium, mature. P, Caryopsis. 


SAARELA ET AL., A NEW SPECIES OF BROMUS FROM PERU 921 


3-6.6 mm long, membranous, pilose adaxially; apex obtuse, erose. Blades 20-55 cm long, 
3-8 mm wide, flat, apically acuminate, mostly sparingly pilose below and glabrous to 
sparingly pilose above with hairs 0.5-1.2 mm long; margins scabrous. Panicles 12-25 cm 
long, 1-8 cm wide, obovate, lax, often nodding; central axis glabrous or with an occa- 
sional hair; branches mostly 1.5-10 cm long, ascending or appressed to spreading and 
drooping, naked near base, often re-branched immediately at base, scabrous, hairs antrorse 
and stiff; lower inflorescence nodes usually with 2-4 branches. Spikelets (12-)18-27 mm 
long, florets 5-8, laterally compressed; lowest rachilla internodes 2-3 mm long. Glumes 
5-11.2 mm long, glabrous or sparsely hairy; apex entire or mucronate, the mucro usually 
< 1mm long; lower glumes 5-8 mm long, l-veined, apex acuminate; upper glumes 8-11.2 
mm. long, 3-veined, apex acute. Lemmas 10-12.6 mm long, chartaceous, 5- or 7-veined, 
pilose, especially along margins and near apex, hairs 0.2-0.5 mm long; margins tightly 
involute on lower 1/5-1/4; apex entire or minutely bifid, teeth < 0.2 mm long, awned, 
awn 3-5.5 mm long, straight. Paleas 9-12 mm long, shorter than lemma, membranous to 
hyaline, ciliate along keel, glabrous between veins; apex acuminate, sometimes mucr- 
onate, veins extending as mucro usually < 1 mm long. Lodicules 1-1.3 mm long, nar- 
rowly elliptic to narrowly obovate, unlobed. Stamens 3; anthers 3.1-3.8 mm long, yellow- 
ish-orange. Ovaries 1.2-2 mm long, golden brown, hairy on upper 1/2-2/3 with hairs 
0.3-0.6 mm long; styles 2, separate; stigmas 2, feathery, whitish. Car yopses 5.8-7 mm long, 
yellowish-brown, apex hairy, hairs 0.4-0.8 mm long; hilum extending length of grain. 

Distribution and Habitat—Known only from near the type locality 12-15 km E of 
Puquio on road towards Cuzco between 3730-3790 m on slopes under large boulders and 
beneath thorny shrubs, such as Colletia spinosissima J. Gmel. (Rhamnaceae), Berberis 
(Berberidaceae), and Ribes (Grossulariaceae); also associated with Lepidophyllum 
(Ast ) and Festuca (Poaceae). Suitable habitat for B. ayacuchensis is estimated to be 
5 km? at the type locality (Fig. 3). 

Etymology—The epithet ‘ayacuchensis is designated to reflect the known distribu- 
tion of the new species in Departamento Ayacucho, Peru. 


Additional Specimens Examined: PERU: Departamento Ayacucho: Provincia Lucanas, 13 km E of Puquio at km 

172 marker (14°41'18.5" S Lat, 74°04'26.8"W Long), 8 Apr 2004, PM. Peterson & N.F Refulio-Rodriguez 18212 (K, 
MO, UBC, US, USM); 15 km E of Puquio at Km 174 — 42.9"S Lat, 74°03'46.8" W Long), 9 Apr 2004, PM. 
Peterson & N.F Refulio-Rodriguez 18233 (kK, MO, UBC, US, USM 


Leaf Anatomy—tThe blades (Fig. 4) are flat and have a C3 anatomy, with completely 
sclerefied margins. The epidermis is comprised of oval to round, subequal cells. Bulliform 
cells are 2-4 times the size of the regular epidermal cells, located adaxially, and arranged 
in groups of 5-6. Stomates are located adaxially and abaxially, and are usually associated 
with deep sunken pits. The chlorenchyma tissue is of the festucoid type. Vascular bundles 
are all primary, round in outline, and they are all approximately the same size, with the 
exception of the larger central rib. Vascular bundles are well differentiated into xylem 
adaxially and phloem abaxially. Vascular sheaths are double; the inner sheath is scleren- 
chymatous and complete, whereas the outer sheath is parenchymatous and incomplete. 
Sclerenchyma fibers are associated adaxially with all bundles, and abaxially with all 
bundles except those towards the margins of the blade; intercostal sclerenchyma absent. 
The central structure of the blade is well differentiated, and includes a vascular bundle 
and a prominent keel comprised of sclerenchyma. 


BRIT.ORG/SIDA 22(2) 


Peru 


250 km 


Fic. 3 hi 


KEY TO THE SPECIES OF BROMUS IN PERU 
Lemma awns geniculate and twisted; plants short-lived annuals (sect. Neobromus) 


Lemma awns straight and not twisted; plants short-lived to long-lived annuals, biennials, or 
P erennials. 


B. berteroanus 
2. Spikelets generally laterally compressed; lemmas laterally compressed and keeled (sect. 
Ceratochloa 
3, Mature anthers 3—5.5 mm lon 

er glumes 1-veined 


upper glumes 3-veined; culms densely pilose below; ligules 
adaxially pilose, 3-6.6 mm long;lemmas pilose, especially along the margins and near 
pex 


naq:lemm 


B. ayacuchensis 
4, Lower glumes 5-veined; upper glumes 7-9-veined; culms glabrous or pubescent be- 
low;ligules adaxially glabrous, 2-3 s glabrous or scabrous throughout 
3. Mature anthers 0.5-2 mm long 
5. 


. cebadilla 
Lemma awns 12—18(-25) cm long; plants short-lived annuals found se ek the 
coastal desert of Arequipa; panicle branches nodding, drooping and flex 


f B. striatus 
5, Lemma awns 0,5-6 mm long; plants long-lived annuals, biennials, or aa plants 


SAARELA ET AL., A NEW SPECIES OF BROMUS FROM PERU 923 


Fic. 4. Leaf anatomy of Bromus paseatcten ss enon eee: nodignen Sh Mansvelse section a A. eu of leaf blade 
Caley mas im); B f blad 75 um 


1 + 
P v t J 7 LUllldald, 2 yer 


more wide-ranging; ee branches usually stiffly erect and ascending, rarely nod- 
ding, drooping, or flexuo 
6. Lower glumes 5-7(-9)- ne upper glumes generally 7—9-veined; spikelets strongly 


laterally compressed B. catharticus 

6. Lower glumes 3-veined; upper glumes 5-veined; spikelets not strongly laterally 
compressed B. coloratus 

2. Spikelets terete or dorsally flattened; lemmas co or dorsally flattened (sect. Bromopsis). 

7. Culms 5—30(—40) cm tall;leaf blades Sia 5mm wide. 

8 Spikelets densely villose; panicles 1-3.5 cm a contracted with short, stiff branches; 
lemmas hairy B. villosissimus 

8. Spikelets glabrous, scabrous or pilose; panicles 3-8 cm long, open with long, flexuous 
branches; lemmas glabrous . modestus 


7. Culms (15-)30-120 cm tall; leaf blades generally 2-12 mm wide. 
9. Lemma awns (4-)5-9 mm long; lower glumes 0.8-1 mm wide near base. 


10. Pedicels flexuous, erect ae as aa deg ee ny B. flexuosus 
10. Pedicels straight and er glabrous 
9, Lemma awns 1—5(-6) mm oe lower glumes ] ay mm ice near a 
11. Lower glumes 1-1.2 mm wide near base; pedicels | hairy; lemmas villous B. lanatus 
11. Lower glumes 2.2-3 mm wide near base; pedicels glabrous to short pubescent; 
lemmas glabrous to pubescent, sometimes villous along the margins B. pitensis 
APPENDIX 1 


Citation of new records and confirmed reports of Bromus in Peru. 


Bromus cebadilla Steud. — Cajamarca: San Miguel:61 km N of Cajamarca on highway 3N towards Bambamarca, 
3640 m, 16 Mar 2000, PM. Peterson 14914 & N.F. Refulio-Rodriquez (ACOR, US, USM). 

Bromus coloratus Steud.—Arequipa. Caraveli: 5 km S of Airoca and 3 km N of Cahuacho, 15°28'10.3"S, 
73°29'26.6'W, 3520 m, 3 Mar 2002, PM. Peterson 16390 & M.I. LaTorre, A. Ramirez & D. Susanibar (ACOR, US, USM). 
Apurimac. Aymaraes: 9 km SW of Cotaruse on road towards Puquio, 3510 m, 14°29'30.8"S, 73°15'04.2"W, 13 
Mar re PM. Peterson 16480 & N.F. Refulio-Rodriguez (ACOR, US, USM). Ayacucho, Lucanas: 6 km E of Puquio on 


924 BRIT.ORG/SIDA 22(2) 


road towards Chavina, 14°41'43.9"S, 74°05'35.4"W, 3274 m, 27 Feb 2002, PM. Peterson 16296 & M1. Lalorre,A. Ramirez 
& D. Susanibar (ACOR, US, USM). Junin. Junin: 2 km SW of San Jose on road towards Junin, 3970 m,8 Apr 1997, 
PM. Peterson 14121 & NF Refulio-Rodriguez (ACOR, USM). 


Bromus flexuous Planchuelo.—Cuzco. Calca: 22 km N of Calca on road towards Lares, 4120 m, 13°12'56.6'S, 
71°54'35.2"W, 16 Mar 2002, PM. Peterson 16556 & N.F Refulio-Rodriquez (ACOR, US, USM). 


Bromus modestus Renvoize.— Arequipa: Sumbay, 4000 m, A.Weberbauer 6902 (F, US). Huancavelica. 
Huancavelica: Tansiri near Manta, 4400-4500 m, 4 Apr 1953, O. Tovar 1181 (US, USM). Ayacucho. Lucanas: 25 
km SE of Puquio on road towards Chavina, 14°45'17.0"S, 74°03'02.6"W, 3850 m, 27 Feb 2002, PM. Peterson 16302 
& MI. LaTorre, A. Ramirez & D. Susanibar (US). Cuzco. Calea: 30 km N of Claca just below Abra de Amparaes, 
13°10'40.2"S, 71°54'09.2"W, 4530 m, 15 Mar 2002, PM. Peterson 16562 & N.F Refulio- Rodriguez (ACOR, US, USM). 
Puno. Chucuito: 20 Km E of Huacullani on road towards Desaquadero, 3880 m,5 Mar 1999, PM. Peterson 14636 
& NE Refulio-Rodriquez, F. Salvador-Perez (ACOR, US, USM). 


ACKNOWLEDGMENTS 


We are grateful to Alice R. Tangerini for providing the illustration; Alain Touwaide for 

correcting the Latin diagnosis; Robert J. Soreng for discussions concerning the new spe- 

cies; Ana Maria Planchuelo for identifying P.M. Peterson collections; Patricia Gomez 

Bustamonte for correcting the Spanish resumen; Sean W. Graham, Neil Snow, Isidoro M. 

Sanchez Vega, and Stephan Hatch for constructive comments on earlier versions of the 

manuscript; and S.WG. for providing funds from his Natural Sciences and Engineering 

Research Council of Canada (NSERC) Discovery grant to support collection of the new 

sequence data. Field collections by PM.P. were supported by the Biodiversity and Inven- 

tory Program and Jose Cuatrecasas Fund (Smithsonian Institution, National Museum of 

Natural History). J. M.S. was supported by scholarships from NSERC, Alberta Ingenuity, 

and the University of British Columbia, and a research grant from the Lawrence R. 

Heckard Endowment Fund of the Jepson Herbarium. 

REFERENCES 

Acepo, C.and F. Liamas. 1999. The genus Bromus L. (Poaceae) in the Iberian Peninsula. Phan. Monog. 
XXII:1-293. 

Ainoucue, M.L. and R.J. Bayer. 1997. On the origins of the tetraploid Bromus species (section Bromus, 
Poaceae): insights from the internal transcribed spacer sequences of nuclear ribosomal DNA. 
Genome 40:730-743. 

Braxo,L.,and J.L.Zarucchi. 1993. Catalogue of the flowering plants and gymnosperms of Peru.Monogr. 
Syst. Bot. Missouri Bot. Gard. 45:1—-1286. 

CAmarA HernaNnoez, J. 1978. Bromus L.|n Correa, M.N., ed. Flora Patagénica. Vol. 8. Colecc. Ci. Inst. Nac. 
Tecnol. Agropecu. Ill. Buenos Aires, Argentina. Pp. 77-93. 

Crayton, W.D. and S.A. Renvoize. 1986. Genera Graminum. Grasses of the World. Kew Bull. Add. Ser. 13: 
1369, 

Cotumeus, J.T. 1999. Morphology and leaf blade anatomy suggest a close relationship between 
Bouteloua aristidoides and B. (Chondrosium) eriopoda (Gramineae: Chloridoideae). Syst. Bot. 23: 


FeiseNstEIN, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 
ee 
Fu, Y.-B., B.E. Coutman, Y.S.N. Ferbinanbez, J. Cavouette, and PM. Peterson. 2005. Genetic diversity of fringed 


brome (Bromus ciliatus) as determined by ie ae length polymorphism. Canad. J. 
Bot. 83:1322-1328. 


SAARELA ET AL., A NEW SPECIES OF BROMUS FROM PERU 925 


Gutierrez, H.F.and J.F. Pensiero. 1998. Sinopsis de las especies Argentinas del género Bromus (Poaceae). 
Darwiniana 35:75-114. 

Hoimstrom, G.and H. ScHotz. 2000.8 bidentat nova and B. lepidus, two odd brome-grasses 
(Gramineae). Bot. Jahrb. Syst. 122:195-200. 

Marrinetto, G.E. and T.T. ScuHirino-Wittman. 2003. Chromosomes of Bromus aleuticus Trin. ex Nees 
(Poaceae). Genet. Mol. Bio. 26:369-371. 

Massa, A.N., S.R. Larson, K.B. Jensen, and D.J. Hote. 2001. AFLP variation in Bromus section Ceratochloa 
germplasm of Patagonia. Crop Sci.41:1609-1616. 

Massa, A.N., K.B. Jensen, S.R. Larson, and D.J. Hote. 2004. Morphological variation in Bromus sect. 
Ceratochloa germplasm of Patagonia. Canad. J. Bot. 82:136-144. 

MattHel, O. 1986. The genus Bromus L. (Poaceae) in Chile. Gayana Bot. 43:47-110. 

Mercatre, C.R. 1960. Bromus. In: Anatomy of Monocotyledons: Gramineae 1.Clarendon Press, Oxford. 

Narano, C.A., FH. Arias, F.E. Git, and A. Soriano. 1990. Bromus pictus of the B. setifolius complex (section 
Pnigma): numerical taxonomy and chromosome evidence for species rank. Canad. J. Bot. 68: 
2493-2500. 

Naranio, C.A. 1992. Estudios biosystematicos en especies de Bromus (seccidn Ceratochloa, Poaceae). 
|. Sistemas reproductivos y barreras de aislamiento. Darwiniana 31:1 73-183. 

Oua, T., V. Jaaska, and V. Vistar. 2003. Breeding system, evolution and taxonomy of Bromus arvensis, B. 
japonicus and B. squarrosus (Poaceae). Pl. Syst. Evol. 242:101-117. 

Pavuck, L.E. 1995. Bromus L.of North America. Royal British Columbia Museum, Victoria. 

Pavuick, L.E., A.M. PLANCHUELO, PM. Peterson, and R.J.Sorenc. 2003. Bromus.Pp. 154—191.|In:Soreng,R.J.,RPM 
Peterson, G. Davidse, E.J. Judziewicz, F.O. Zuloaga, IS. Filgueiras,and O. Morrone, Catalogue of New 
World Grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1-730. 

Peterson, PM. and A.M. PLancHueLo. 1998. Bromus catharticus in South America (Poaceae: Bromeae). 
Novon 8:53-60. 

Peterson, PM., J. Cayouette, Y.S.N. FERDINANDEZ, B. CouLMAN, and R.E. CHapman, 2002. Recognition of Bromus 
richardsonii and B.ciliatus: evidence from morphology, cytology, and DNA fingerprinting (Poaceae: 
Bromeae). Aliso 20:21-36. 

Pitay, M.and K.W. Hitu. 1990. Chloroplast DNA variation in diploid and polyploid species of Bromus 
(Poaceae) subgenera Festucaria and Ceratochloa. Theor. Appl. Genet. 80:326-332. 

Pictay, M.and K.W. Hiv. 1995.Chloroplast-DNA restriction site analysis in the genus Bromus (Poaceae). 
Amer. J. Bot. 82: 239-249. 

Pinto-Escosar, P. 1981.The genus Bromus in northern South America. Bot. Jahrb. Syst. 102:445-457. 

Pinto-Escosar, P1986. El genero Bromus en Los Andes Centrales de Suramerica. Caldasia 15:15-34. 

PLANCHUELO, A.M. 1983.Una nueva especie de Bromus (Poaceae) de Argentina. Kurtziana 16:123-131. 

PLancHueto, A.M. 1991.Estudios sobre el complejo Bromus catharticus (Poaceae) |.Evaluacion estadistica 
de los caradcteres taxonomicos. Kurtziana 21:243-257. 

PLANCHUELO, A.M.and P.M. Peterson. 2000.The species of Bromus (Poaceae: Bromeae) in South America. 
In: Jacobs, S.W.L. and J. Everett, eds. Grasses: systematics and evolution. CSIRO, Melbourne. Pp. 
89-101. 

Ramos, J.C., J.C. Tivano, and A.C.Vecetti, A.C. 2002. Anatomical studies of reproductive stems of Bromus 
qauleticus Trin. ex Nees. (Poaceae). Gayana Bot. 59:51-60. 

Renvoize, S.A. 1994. Notes on Sporobolus and Bromus (Gramineae) from the Andes. Kew Bull. 49: 
543-546. 

Renvoize, S.A. 1998. Gramineas de Bolivia. Royal Botanic Gardens, Kew. 

SAARELA, J.M., PM. Peterson, R.M. Keane, J. Cavouette, and S.W. Granam.|In Press. Molecular systematics of 


926 BRIT.ORG/SIDA 22(2) 
Bromus (Poaceae: Pooideae) based on chloroplast and nuclear DNA sequence data.In: Columbus, 
J.T.,E.A. Friar, J.M. Porter, L.M. Prince, and M.G. Simpson, eds. Monocots: comparative biology and 
evolution, 2 vols. Rancho Santa Ana Botanic Garden, Claremont, California, USA. 

SAARELA, J.M., RBM. Peterson, and J. Cavouette. 2005. Bromus hallii (Poaceae), a new combination for Cali- 
fornia, and taxonomic notes on B. grandis and B. orcuttianus. Sida 21:1997-2013. 

SmitH, P.1970.Taxonomy and nomenclature of the brome-grasses (Bromus L.s..). Notes Roy.Bot. Gard. 
Edinburgh 30:361-375. 

SoperstroM, T.R.and J.H. Beaman. 1968. The genus Bromus (Gramineae) in Mexico and Central America. 
Publ. Mus. Michigan State Univ. Biol. Ser. 3:465-520. 

Stessins, G.L. 1981.Chromosomes and evolution in the genus Bromus (Gramineae). Bot. Jahrb. Syst. 
102:359-379. 

Tovar, O. 1993. Las Graminees (Poaceae) del Peru. Ruizia 13:1-480. 

Tuna, M., K.P. Vocet, and K. ARUMUGANATHAN. 2005. Genome size and Giemsa C-banded karyotype of 
tetraploid Bromus ciliatus L. Euphytica 146:177-182. 

Tuna, M., K.P. Vocet, and K. ARUMUGANATHAN. 2006. Cytogenetic and nuclear DNA content characteriza- 
tion of diploid Bromus erectus and Bromus variegates. Crop Sci.46:637-641. 

Waanon, H.K. 1952. A revision of the genus Bromus, section Bromopsis, of North America. Brittonia 

7415-480. 


PSIDIUM CAULIFLORUM (MYRTACEAE), 
A NEW SPECIES FROM BAHIA, BRAZIL 


Leslie R.Landrum Marcos Sobral 
School of Life Sciences Departamento de Botanica UFMG 
Arizona se University Caixa Postal 486, 31270-901 
Tempe, AZ 85287-4501, U.S.A. Belo Horizonte MG, BRASIL 
les. pares sobral@ufmg.br 
ABSTRACT 


Psidium cauliflorum, from the Brazilian state of Bahia, is described and illustrated. It differs from all other spe- 


Ider branches and trunks. It may belong toa complex 


cies of Psidium in having its flowers borne in cht 
of species ane P. oligospermum, P. sartorianum, P. glaziovianum, P. schenckianum, and perhaps P. 
appendiculatum 


RESUMO 


Psidium canlilonains uma nova ESDEGIe do este ea Bahia, Brasil, é descrita e ilustrada. Essa espécie, distinta das 
log p ramos vento e ieneos é possivelmenite ESlacionada ao 


1 inclui Poli Dea esa Pel 
a L Oo L Oo yo ig 


Psidium L. (Myrtaceae) is a genus of at least 50 and perhaps as many as 100 species 
(McVaugh 1968) with a natural range from Mexico and the Caribbean to Uruguay and 
northern Argentina on the American continents and extending to some east Pacific is- 
lands (e.g., Galapagos). Psidium is distinguished from other genera by a combination of 
floral and seed characters discussed in Landrum and Sharp (1989) and Landrum (2003). 
Psidium cauliflorum appears to belong to a group of species including P oligospermum 
DC,, P. sartorianum (O. Berg) Niedenzu, P glaziovianum Kiaerskou, P. schenckianum 
Kiaerskou, and perhaps P. appendiculatum Kiaerskou. These species have 
brochidodromous leaf venation with dendritic tertiaries. The calyx is generally closed 
or nearly closed in the flower bud and they often have wart-like to flange-like apical 
appendages on the calyx (but this character is not present in P. cauliflorum). The seeds 
are 3-6 mm long with rounded edges and few (5-12) per fruit. Ovules are usually less 
than 30 per locule. The limits between widespread and variable P sartorianum and P 
oligospermum, which is restricted to Bahia, are still problematic and require more study. 
We believe that Psidium oligospermum and P schenckianum frequently hybridize based 
on herbarium specimens and field observations. All of these species (except perhaps P 
sartorianum) grow in Bahia and are sometimes endemic to that state. 


Psidium cauliflorum Landrum & Sobral, sp. nov. (Fig. 1). Typr: BRAZIL. Barta: mun. Cachoeira, 
aes Bee, Vale dos Rios Paraguacu e Jacuipe, 39°05'W, 12°32'S, 40-100 m, Dec 1980 (fl), Grupo Pedra do 
CH E: HRB, =ASU, BHCB photos; isotypes: ALCB (2 sheets),= ASU, BHCB photos, RB). 


Haec species Psidio sartoriano primo aspectu similis, flores bascieulat seca even ee 3-5.5- mietrals) 


folia elliptica, vel ovalia, 2.7-6.8 x 1-3 cm, 1.8-2.6 plo longiora quam | 
clausus, sub anthesi irregulariter fissus. 

Tree 3-5.5 m high, sparsely to densely pubescent on young growth; hairs rusty brown to 
whitish, mostly erect or spreading, up to ca. 1 mm long; young twigs terete to slightly 
compressed, densely to moderately pubescent, the bark reddish brown to gray, the older 


SIDA 22(2): 927 — 929. 2006 


928 


= 


BRIT.ORG/SIDA 22(2) 


AS anit Li ad 
am Sa Sigh la 
; yp vig 
Fis. 1. A- B, De? 43 Ul A L 4 | R "4 fei 1 | a a 141 twill D poionne of 
ee an two ane buds E. detached hab : anthers, G. remnants of flowers and young fruit after 
alle y ation, Lf | | d.(A & 1, Grupo ‘peda 


Pedra do Cavalo 955 /HRB, the holotype; D-H, Queiroz et al. 1742, ASU). Drawn by Bobbi Angell. 


do Cavalo oe ALCB; C, G 


RAITT DTILATALITCADD Ai 929 


twigs gray, glabrous, the bark somewhat flaky. Leaves elliptic to oval, 2.7-6.8 cm long, 1- 
3 cm wide, 1.8-2.6 times as long as wide, moderately to sparsely puberulent (or more 
densely so along midvein), glabrescent with age; apex acute; base rounded to cuneate: 
petiole channeled, densely to sparsely pubescent to glabrescent, 2-3 mm long, 0.5-1 mm 
wide; midvein impressed proximally to nearly flat distally above, prominent below, the 
venation brochidodromous, the lateral veins slightly raised and visible above or obscure, 
6-9 pairs leaving midvein at an angle of ca. 45 degrees, the marginal vein arching be- 
tween laterals, equalling them in prominence, running ca. 1-1.5(-3) mm from margin, the 
tertiary veins forming a dendritic pattern that arises from the marginal vein, scarcely to 
clearly visible; blades subcoriaceous, drying reddish brown to grayish, densely glandular 
beneath. Flower buds pyriform, 4-6 mm long, densely pubescent on hypanthium, sparsely 
so on calyx, borne on older stems in clusters of as many as 20, apparently appearing at 
the same point season after season; peduncles uniflorous to triflorous, 1-9 mm long, ca. 
0.6 mm wide, sparsely to densely pubescent, sometimes borne on short bracteate shoots, 
the branches of dichasia ca. 3 mm long; bracteoles linear to narrowly lanceolate, ca. 1-15 
mm long, caducous before anthesis; calyx closed except for an apical pore, with hairs 
sometimes protruding from pore, tearing irregularly at anthesis; petals obovate to subor- 
bicular, ca. 7 mm long, glabrous or with cilate margins, hypanthium obconic, 2-3 mm 
long; disk ca. 3.5 mm across at anthesis (5 mm in fruit), pubescent; stamens ca. 6 mm 
long, 150-190; anthers ca. 0.3mm long, witha terminal gland and usually 2 other glands 
in the connective; style ca. 8 mm long, glabrous; ovary 3-locular; ovules 7-20 per locule, 
uniseriate on each lamella of a slightly peltate placenta. Fruit 1-2 cm in diam.; seeds few, 
ca.6mm long. 

Only five collections have been made of Psidium cauliflorum so little is known of 
this very distinctive, but rare species. It has been collected flowering in June, July, Octo- 
ber, and December and probably fruits shortly afterward as is normally the case in 
Psidium. De Queiroz et al. report that it grows in “floresta estacional,” that is, temporarily 
deciduous dry forests. 


Paratyres: BRAZIL. Banta: mun. Cachoeira, Estacgao da Mata, Vale dos Rios Paraguacu e Jacuipe, 39°05'W, 12°32'S, 
40-120 m, Jul 1980 (£1, fr), Grupo Pedra do Cavalo 407(ALCB), Jun ie a fr), Grupo Pedra do Cavalo 326 (ALCB), 
Oct 1980 (£D, Grupo Pedra do Cavalo 814 (ALCB, CEPEC); mun. Feira de Santana, 11 km NW de Jaguara, Fazenda 
Monte Verde, floresta estacional, 320 m, 21 Jul 1987 (£1), L. P de Queiroz, Lemos & Lébo 1742 (ASU). 


ACKNOWLEDGMENTS 


We thank the curators of the following herbaria for allowing us to study their specimens 
for the preparation of this paper (ALCB, ASU, CEPEC, HRB) or for sending specimens in 
exchange (HUEFS). Travel in Brazil was made possible for LRL through an American 
Republics Fulbright Grant. We thank Carolyn Proenca and Fred Barrie for helpful re- 
views of this manuscript. 


REFERENCES 


LanoruM, L.R. 2003. A revision of the Psidium salutare complex (Myrtaceae). Sida 20:1449-1469. 

Lanorum, L.R. and W.P. SHare. 1989. Seed coat characters of some American Myrtinae (Myrtaceae): 
Psidium and related genera. Syst. Bot. 14:370-376. 

McVauch, R. 1968.The genera of American Myrtaceae — an interim report. Taxon 17:354-418. 


BRIT.ORG/SIDA 22(2) 
Book REVIEW 
Cuaries Ainsworth (Ed.). 2006. Annual Plant Reviews, Volume 20: Flowering and Its Ma- 
nipulation. (ISBN 1-4051-2808-9, hbk.). Blackwell Publishing, 9600 Garsington Road, 
Oxford OX4 2DQ, UK. (Orders: www. blackwellpublishing.com, 1-800-862-6057). 
$199.99, 304 pp., 61/4" x 91/2" 


Anastonishing variety of flowering plants currently dominate the plant life of our planet. The editor of Flower 
ing and Its Manipulation, Charles Ainsworth, writes in the preface“ 


there are good reasons for supposing that the 
flower itself has been a ms contributing factor to the spread of the Angiosperms.” 


mh 


Therefore, the study of 
giosperm flowering is an important field. This genetics-heavy volume from the Annual Plant Reviews serie 
designed to complement an earlier edition- Volume 6: Plant Reproduction. The chapters of the book are laid out in 
a logical order that parallels the birth, development, and death of the flower. The first chapter is devoted to a 
discussion of the origin and evolution of the flower through a genetic model. Chapter 2 focuses on the study of 
floral induction, or the factors that cause plants to flower at certain moments in time. The goal of Chapter 3 is to 
“outline the present state of ee of on unction, regulation, and conservation of the genes that partici- 
pate in the ABC program...” the ddel 


an- 


oral development. The complex genetic lees that underlie 
the beguilingly simple ABC model are clearly ae In chapter 4, the “genetic control of flower size and 
shape” is discussed in detail. The next chapter focuses on Ty peuetics and development of floral architecture, de- 
scribed as “.the shoot system that gives rise to flow 


— 


The second part of the book includes three chapters 
focused on “specialized components of floral selon Chapter : mee on tl 


he complex patterns of male 
and female [lowers in monoecious plants, with a focus on the well-st ucumber and maize plants. Chapter 


he diversity and ee of perennials is the focus of 
Chapter 8, which provides “a sketch map of the types of perennial flowering.” The next two chapters are devoted 
to the most celebrated floral aspects- color and scent. Chapter rg acta: recent edeveloprent:) in our understand- 
ing of the biochemistry and molecular biology of flower color. seq hapter describes the biochemis 
try, physiology, genetics, and regulation of floral scent. Both chapt tain details on the use of genetic engi- 
neering to manipulate floral color and scent. The final chapter is “appropriately” focused on flora 
according to the editor. Floral senescence, or the “death” of f 


Eu focuses on the genetics ol f cytoplasm ic male aie T 


—_ 


— 


senescence, 
‘lowers, is discussed in detail, as well as methods used 
to manipulate senescence and useful applications for the floral industry.—Marissa Oppel, MS, Collections and Re- 
search Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A. 


SIDA 22(2): 930. 2006 


NEW SPECIES OF MYRTACEAE FROM ECUADOR AND PERU 


Bruce K. Holst Maria Lucia Kawasaki 
Marie Selby Botanical Gardens Botany - The Field Museum 
811 South Palm Avenue 1400 south Lake Shore Drive 
Sarasota, Florida 34236, U.S.A. Chicago, Illinois 60605-2496, U.S.A. 
bholst@selby.org lkawasaki@fmnh.org 
ABSTRACT 


Two new species of Myrtaceae from Ecuador and Peru are described and illustrated: Calyptranthes manuensis 
and Eugenia yasuniana 


RESUMEN 


+7 ] s 4Jokf ] 


1 Ecuador y P 


Ged +] 7 = a 
VEUCSLIITVEOLL ¥} Df EUStilla 


yasuniana. 

INTRODUCTION 
In preparation for the treatment of Myrtaceae for the Flora of Ecuador and adding to the 
checklist of species for the country (Holst 1999), two new species are described in this 
paper: Calyptranthes manuensis and Eugenia yasuniana. 


Calyptranthes manuensis sD. Holst & M_L. Kawasaki, sp. nov. (Figs. 1 1A—G). Type: PERU. MADRE 
DE Dios: Manu, P 1 Manu, Rio Manu, Cocha Cashu Station, 350 m, 8 Oct 1980 (f]), R. Foster 
5501 (HOLOTYPE: F; ISOTYPE: SEL). 


Ab omnibus speciebus generis characteribus combinatis differt: inflorescentiis reductis, alabastris oblongis, 
fAl14 


subsessilibus vo medio supra biconvexo. 


Shrubs or small trees 1-10 m high, mostly glabrous except for the sericeous young veg- 
etative growth, trichomes dibrachiate, yellowish-brown; branchlets terete, often with l 
or 2 pairs of narrowly elliptic, navicular bracts, 1-2.5 x 0.2-0.4 cm. Young leaves droop- 
ing, reddish; leaf blades narrowly to broadly elliptic to narrowly obovate, 14-18 x (3-)5- 
8 cm, chartaceous, thinly sericeous when very young on lower surface, becoming mostly 
glabrous, drying brownish-green, paler below; apex abruptly acuminate, the acumen to 
ca. 1 cm long: base cuneate to obtuse; midvein biconvex above, convex below; lateral veins 
20-25 pairs, slender, inconspicuous, slightly convex on both surfaces; marginal veins 2, 
the innermost 2-3 mm from blade margin, similar to the lateral veins; glands numerous, 
light-brown, salient on both surfaces; petiole 0.5-1.2 cm long, flattened, glabrous. Inflo- 
rescences paired, greatly abbreviated, 2-flowered spikes borne at the nodes of the fallen 
bracts, ca. 1.5-2 cm long, the main axis 2-3 mm long, densely sericeous to glabrous, the 
pedicels 0-5 mm long, stout when present. Flower buds oblong, 8-10 mm long, gland- 
dotted; bracteoles subulate, ca. 2mm long, early deciduous; calyx calyptrate, puberulous 
to glabrous, acuminate, deciduous; petals absent; stamens numerous, the filaments ca. 4- 
5 mm long, the anthers ca. 0.5 mm long; style ca. 6 mm long, the stigma capitate; hy- 
panthium prolonged ca. 3-4 mm beyond the ovary, the upper portion explanate at an- 
thesis, densely puberulous basally to glabrous without; disk ca. 3 mm diam.; ovary 
2-locular, with 2 ovules per locule. Fruits globose, 2-3 cm diam., crowned by a circular 
hypanthium scar, gland-dotted, glabrous, dark-purple to black; seeds 1 or 2,ca.9 x 7mm, 
the seed coat membranous; embryo myrcioid, the cotyledons leafy and folded, the radicle 
well developed, equaling cotyledons in length. 

Distribution —This species, first known from Amazonian Peru (Parque Nacional del 


SIDA 22(2): 931 — 934. 2006 


932 BRIT.ORG/SIDA 22(2) 


Scale = 1 cm 


D 


Fic. 1. G lyy tranth is B. Holst & M.L. Kawasaki. A. l | fruits. B. Calyptra C. Flower. D. Ti ti 
ovary. E. Fruits. F. Detail of infl G. Detail of | hl ith icular t (A, Foster et al. 7186; B—D, F, Foster 5507; E, Foster 
& Terborgh 5244; G, Croat 72503). 


Manu in Madre de Dios), was later collected in Ecuador; it occurs in lowland humid (flood- 
plain) forests, at 320-350 m elevation. 

Calyptranthes manuensis is characterized by the much reduced inflorescences, 
subsessile to short-pedicellate, oblong flower buds, and leaves with biconvex midvein. 


Additional collections examined: ECUADOR. Napo: Jatun Sacha, Rio Napo, at mouth of Rio Huambuno, 3 km 
downstream from Campana Cocha ane forest on river floodplain, 00°55'S, us 25'W, 350 m, 29 Jun 1987 (fr), D. 
Neill, W. Wilbert & SFS Students 7742 (MO, SEL). Pastaza: Via Auca, 115 km S of Coca, ca. of Rio cur road 
PETRO- CANADA in nstruction, 01°] a se 55'W, 320 m, 24 May 1989 (fr), : aa 33 (FR MO, SEL): V 
115 km S of Coca, ca. of Rio Tigtino, road PETRO-CANADA in construction, 01°15'S, 76°55'W, 320 m, 24 io ie 89 
(fr), D. Rubio 153 (F MO, SEL); Via Auca, 110 km S of Coca, 10 km from ae Tigtiino, Sector Cristal, 01°15'S, 76°55'W, 
320 m, 7 Jan 1989 (fr), W. Palacios, C. Iguago & F Hurtado 3361 (F, MO, SEL). Sucumbios: Lago Agrio, along road 
between Lago Agrio (Nueva Loja) and Coca (Pto. Francisco de ane 26 km S of Lago Agrio, 4.6 km S$ of El 
Emo, then 2.8 km W of main Lago Agrio-Coca Road, along farm road, 00°05'S, 76°54'W, 355 m, 29 Feb 1992 (fr), T. 
Croat 72503 (MO, SEL). PERU. Madre de Dios: Manu, Parque Nacional del Manu, Rio Manu, vicinity of Cocha 
Cashu Station, 5 Nov 1976 (fD, R. Foster & J. Terborgh 5189 (F); Manu, Parque Nacional del Manu, Rio Manu, vicin- 
ity of Cocha Cashu Station, 27 Nov 1976 (fr), R. Foster & J. Terborgh 5244 (F); Manu, Parque Nacional del Manu 
Rio Manu, vicinity of Cocha Cashu Station, 22 Apr 1977 (fr), R. Foster & C. Janson 6279 (F); Manu, ee aciorial 
del Manu, Rio Manu, Cocha Cashu Station, 22 Oct 1979 (y fr), R. Foster 7186 (F); Manu, Parque Nacional del Man 


HOLST AND KAWASAKI, NEW SPECIES OF MYRTACEAE FROM ECUADOR AND PERU 


Fic. 2. E 9 j j B. Holst & M.L. Kawasaki. A. Leaves. B. Inflorescence. C.F 


| 
/ 


lohes have fallen. D, 


Medial section of fruit. E. | d (A—C, E, Romoleroux 2021; D, Villa & Vélez 852). 


934 BRIT.ORG/SIDA 22(2) 


Rio Manu, Cocha Cashu Station, 4 May 1981] (fr), R. Foster & C. Janson 8385 (F); Manu, Parque Nacional del Manu, 
Rio Manu, Cocha Cashu Station, 11°50'S, 71°25'W, 350 m, ie 1984 (fr), R. Foster a (F MO). 


Eugenia yasuniana B. Holst @ M.L. Kawasaki, sp. nov. (Figs. 2A-E). Typr: ECUADOR. ORFLLANA 
NAPO” on label): Estacion Cientifica Yasuni, Tiputini River, NW of confluence with Tivacuno River; 6 

Ga Eof main Maxus Road, Km 44, on spur road toTivacuno oilwell, near 50 ha plot, sendero Norte, 00°38'S, 
76°30'W, 200-300 m, 21 Nov 1995 (fD, K.R & R. Foster 2021 (HOLOTYPE: QCA; ISOTYPES: F SEL). 


NA ‘ Rs oats ae ; | } a iY a dials rere eee | 


es, 


Oo oa fo] 
praesenti (nec absenti), pedicellis brevioribus, calycis lobis deciduis (nec persistentibus) et fructibus globosis 


(nec ellipsoideis). 
Trees 10-12 m high, the trichomes simple, yellowish-brown to yellowish on leaves, red- 
dish-brown to yellowish-brown on inflorescences; branchlets slightly compressed in cross 
section. Leaf blades narrowly elliptic to elliptic, 15.5-23 3 7.5-14 cm, coriaceous, the up- 
per surface puberul abrous, drying brownish-green to dark-brown, the lower sur- 
face paler, densely sericeous-pubescent, glabrescent; apex abruptly acuminate, the acu- 
men to ca. | cm long; base cuneate to obtuse; midvein narrowly convex, in a channel, 
appearing sulcate above, convex below; lateral veins 10-13 pairs, impressed to slightly 
convex above, convex below, arcuate-ascending, the arches forming a marginal vein to 5 
mm from blade margin; glands indistinct to numerous, punctiform, dark-brown, plane 
to convex above, indistinct below; petiole ca. 2 cm long, channeled, puberulous to gla- 
brous. Inflorescences racemose, sometimes shortly so and appearing fasciculate, borne 
on leafless branches, with up to ca. 8 flowers, the main axis to 4 cm long, the pedicels to 1 
cm long, appressed-pubescent. Flowers 4-merous; buds subglobose, ca. 1.5m long: bracts 
and bracteoles suborbicular, ca. 3mm long, persistent; calyx-lobes ovate to broadly ovate, 
ca. 1 cm long, obtuse, sericeous-pubescent without, deciduous; petals elliptic, ca. 1 cm 
long; stamens numerous, the filaments ca. 6 mm long, the anthers ca. 1.2 mm long: style 
ca. 7mm long, the stigma punctiform; hypanthium not prolonged beyond the ovary, ap- 
pressed-pubescent without; disk ca. 4-5 mm diam., pubescent; ovary 2-locular, with sev- 
eral ovules per locule. Fruits globose, ca. 3-4 cm diam., appressed-puberulous, yellow; 
seeds 1-2, ca. 2.5 31.5cm long, the seed coat membranous; embryo eugenioid, the cotyle- 
dons fleshy, connate, the radicle indistinct. 

Distribution.—Known only from Ecuador, in lowland forests of the Estacion 
Cientifica Yasuni (Orellana), at 200-300 m elevation. 

nid yasuniana is related to E. myrobalana DC., a species from Amazonian Peru 

and Brazil, but differs by the leaves being densely sericeous-pubescent below, with a mar- 
ginal vein (vs. leaves puberulous below, marginal vein absent), pedicels to 1 cm long (vs. 
L225 rong? deciduous (vs. persistent) calyx-lobes, and globose (vs. ellipsoid) fruits. 


Additional coll |: ECUADOR. Orellana: Estacion Cientifica Yasuni, Tiputini River, NW of conflu- 
ence with Tivacuno River, 6 km E of main Maxus Road, Km 44, on spur road toTivacuno oilwell, 50 ha plot, 
sendero Saino, 00°59’S, 77°45°W, 200-300 m, 6 ies 1999 (fr), : Romoleroux, G. Villa & L. Vélez 3210 (F QCA): 
Estacion Cientifica Yasuni, Tiputini River, NW of confluence with Tivacuno River, 6 km E of Repsol-YPF Road, 
Km 7, on spur road to oilwell, 00°38'S, 76°30"W, 200-300 m, 29 Jan 200] (fr), G. Villa & L. Vélez 852 (F QCA). 


ACKNOWLEDGMENTS 
We thank Robin Foster and the herbaria cited in the manuscript for the collections. Fred 
Barrie and an anonymous reviewer offered suggestions to help improve this manuscript. 
REFERENCES 


Hovst, B. 1999. Myrtaceae. In PM. Jorgensen and S.Leén-Yanez, eds. Catalogue of the vascular plants 
of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:618-622. 


ZEUXINE PANTLINGII SP. NOV.(ORCHIDACEAE), 
A NEW SPECIES FROM INDIA 


Avishek Bhattacharjee! and H.J.Chowdhery 


pact 


= 


Central Nationa Botanical Survey of India 
Botanical ( IU der , Howrah- 711 103, INDIA 


ABSTRACT 


A new species of Zeuxine Lindl. Zeuxine pantlingii Av. Bhattacharjee @ HJ. Chowdhery closely allied to Z. 
goodyeroides Lindl. is described from West Bengal, India. A brief life sketch and a portrait of Robert Pantling after 
whom the species is named is also provided. 


Key Worps: Orchidaceae, New species, Zeuxine pantlingii, West Bengal, India 


RESUMEN 
Se essen pe una nueva esp cle de Nest Bengal, India de Zeuxine Lind, Zeuxine pantlingii Av. BI harjee @ HJ 
Chowdhery n Z. goodyeroides Lindl. S bién un breve resumen de la vida y 


un retrato a Robert Pantling a quien se dedica la especie. 


Dressler (1993) placed the genus Zeuxine Lindl. in the subtribe Goodyerinae Klotzsch 
under the tribe Cranichideae Endlicher of the subfamily Spiranthoideae Dressler. It is 
represented by about 70 species (Pearce & Cribb 2002) widely distributed from tropical 
Africa, Madagascar through India, Sri Lanka and to the Pacific Islands while, 16 species 
are known from India (Singh et al. 2002). During a recent plant collection trip to North 
Bengal in connection with the revisionary studies of the family Orchidaceae for Flora of 
India project of Botanical Survey of India, the authors came across an interesting speci- 
men of Zeuxine Lindl. A critical study of the literature and herbarium material revealed 
it to be a hitherto unknown species of the genus Zeuxine, which is described and illus- 
trated here. 


Lene panes Av. Bhattacharjee @ HJ. Chowdhery, sp. nov. (Figs. 1-3). Type: INDIA. WEST 
GAL: Darjeeling District, near Mongpoo, ca. 900 m, 28 Aug 2005, A. Bhattacharjee 34807 (HOLOTYPE 
ae 


Zeuxini goodyeroidi Lindl. arte affinis, sed foliis non albocinereo-coloratis; bracteis dorse purpureoviridibus; 
infloresentiis laxis, 2-3(-4) floriferis; floril ibus; hypo ene pee ae epichilis 
mesochilis plus minusve ad 90°, pollinario tension. multo longistipato et 


Terrestrial, rhizomatous herb, 14.5-24 cm long. Rhizome subterranean, creeping, hardly 
distinguishable from the aerial stem, rooting at nodes, 3-5 cm long. Roots dull yellow- 
white, fleshy, tomentose, 3-6 cm long. Aerial stem erect, terete, glabrous, purple-green, 3- 
6cm long. Leaves 4-6, petiolate; petiole sheathing at base, sheath membranous, clasping, 
pink-white, 7-10 mm long; lamina obliquely ovate-lanceolate, entire, acuminate, glossy, 
with dark velvety-green upper surface and purple-green lower surface, 12-36 « 7-15 mm, 
7-veined. Inflorescence 2-3(-4) flowered in very lax spike. Spike terminal, peduncle ter- 
ete, sparsely tomentose, 7-10 cm long, with 2 distant sheaths; sheaths ovate, cuspidate, 
clasping, membranous, glabrous, pink, 8-13 x 2-3 mm, 3-veined; rachis terete, sparsely 
tomentose, l.8-3 cm long. Flowers resupinate, 1.2-1.6 cm long, pink-white, not fully open- 


' Corresponding author's email: avibsi@rediffmail.com 


SIDA 22(2): 935- 940. 2006 


BRIT.ORG/SIDA 22(2) 


ok 


Ne “\ 
WY mm 


e 


Fic. 1. Zeuxine pantlingii Av. Bhattacharjee & H.J. Chowdhery; a. habit, b. entire plant, c. flower, d. bract, e. sepals and petals, f. lip (top 
view), g. lip (lateral view) with bifid rostellum, h. column (in bud), i. anther cap, j. pollinia with stipe and viscidium (front), k. pollinia 
with stipe (tegula) and viscidium (back). 


Fic. 2. Perig (di 1) of Zeuxine | lingii Av. Bk harjee & H.J. Chowdhery 


ing. Floral bracts triangular-ovate, abruptly acuminate, more or less equaling the ovary, 
sparsely pubescent dorsally at basal region, finely serrated at margins, purplish-green, 
9-11 x 4-4.5 mm (at widest end), 3-veined. Sepals unequal, free, glabrous; dorsal sepal 
ovate, acute, forming a hood with petals, pink-red, +.5-5.5 « 2-3 mm, |-veined; lateral 
sepals narrowly lanceolate, sub-acute, proximally dull green, distally pink-red, 5-6 = 1-2 
min, l-veined. Petals obliquely falcate, acute, thin, with scattered minute dots, apically 
connate at a point, glabrous, white, 3-4 « 1-1.5 cm, l- veined. Lip connate to the column 
base, white with light pink tinge, 4-5 mm long; hypochile broadly saccate, 2-3 mm broad 
when spread, each side internally with 1 erect, cornute, ca. 1 mm long lateral gland or 
appendage; mesochile very short, margins incurved to form a flap like structure on both 
sides, flap with minute hairs; epichile simple, obtuse, more or less perpendicular to 
mesochile, surface finely granular, 1-1.5 mm in diameter, margins slightly incurved. Col- 
umn ca. 1 mm high, weakly winged below. Anther beaked, erect; anther cap narrowly tri- 
angular, dull white with red tinge, 3-3.5 mm long; pollinia 2, sectile, clavate, clefted, dull 
yellow-white, ca. 4 mm long; caudicles short; stipe (tegula) linear, semi-transparent, ca. 
2.5mm long; viscidium thick, oblong, dull straw-yellow. Rostellum bifid, 2-2.5 mm long; 
stigma with 2 lateral lobes, ovate; ovary pale-green, 11-12 « 2-2.5 mm. 

Closely allied to Zeuxine goodyeroides Lindl. but differs in having leaves without 
any white-gray midvein; more broader, dorsally purplish-green bracts; larger flowers, 2-3 
(-4) in lax inflorescence; hypochile with longer lateral appendages, epichile more or less 


938 BRIT.ORG/SIDA 22(2) 


: sige tie Pa Meri 9 f 
B.S ee he OR 


Spo ES: 
re 


a 
gé. 


PAR 
cane 


Fic. 2. Poll . £ 7, 7 . sy 17, 


perpendicular to the mesochile; longer pollinarium with much longer stipe and very thick, 
dull straw-yellow viscidium. 

Flowering —August-September 

Distribution.—India, West Bengal, Darjeeling District, near Mongpoo, ca. 900 m. 

Zeuxine pantlingii Av. Bhattacharjee @ HJ. Chowdhery is closely allied to Zeuxine 
goodyeroides LindL, but can be distinguished from several characters in Table 1. 

Etymology.—The new species is named in honour of Robert Pantling (1856-1910), a 
British botanist known for his masterly drawings and colour paintings of Indian orchids. 

Robert Pantling (1856-1910) (Fig. +) was borne in North of England in the year 1856. 
His interest and passion for flowers helped him to get an apprenticeship in Duke's ser- 
vice at the age of 16. After three year’s apprenticeship at the age of 19, Robert Pantling 
was admitted to the Royal Botanic Garden on 5 July, 1875. He left Kew on 14 July 1877 to 
join the Royal Botanic Gardens at Windsor. He came to India in 1879 as an assistant in 
Cinchona Department and subsequently appointed as Curator in the Royal Botanic Gar- 
den, Calcutta, but his deteriorating health and a new post of Senior Assistant took him 
back to Mongpoo in the Eastern Himalayan foothills in Darjeeling district where he spent 
his remaining service in India. At Mongpoo apart from carrying out official duties Pantling 
undertook many plant collection tours to Sikkim Himalayan region and collected hun- 
dreds of orchids. He prepared their drawings and published several new genera and a 


Cher Ae 


Fic. 4. Painting of Robert Pantling (1856-1910). 


Tase 1. Comparison between Zeuxine goodyeroides and Zeuxine pantlingii. 


Z. goodyeroides Lindl. 


Z. pantlingii Av. Bhattacharjee & H.J. Chowdhery 


Leaves with a distinct, white-gray midvein. 
Inflorescence 3-16 flowered in comparatively dense 


spike. 
Bract narrower at base. 


Hypochile with shorter lateral glands or appendages 
within. 

Epichile more or less horizontal to mesochile. 
Pollinarium shorter, with shorter stipe (tegula). 
Viscidium thin, transparent (in live specimen). 

Stipe slightly constricted near the middle (Fig. 3). 


ves without any white-gray midve 
see 2-3(-4) flowered in very uf spike. 


Bract broader at base. 


Hypochile with longer lateral glands or appendages 
within. 

Epichile more or less perpendicular to mesochile. 
Pollinarium longer, with longer stipe (tegula).Viscidium 
very thick, dull straw-yellow (in live specimen). Stipe 
linear (Fig. 3). 


940 BRIT.ORG/SIDA 22(2) 


very large number of new species in collaboration with George King, the then in-charge 
of Calcutta Botanic Garden. His hard official duties, field work and studying minute flo- 
ral details of orchids and drawing large number of orchid illustrations impaired his eye- 
sight and further deteriorated his health. Pantling’s ailing health did not permit him to 
publish any work on orchids after his classic work on orchids, “Orchids of Sikkim 
Himalaya” in 1898, co-authored with George King. Pantling died at the age of 54 at Suez 
on his way back home. He served in India for 31 years. He was commemorated by the 
orchid genus Pantlingia and a large number of species. 

Robert Pantling will be known for his meticulous, beautiful and unmatched, highly 
accurate scientific drawings and paintings of orchids of India, Myanmar and Malaysia 
drawn by him with great precision, which shows his skill and mastery over the subject. 
These drawings are preserved in the archival section of Central National Herbarium, 
Botanical Survey of India, Howrah. 


ACKNOWLEDGMENTS 


The authors wish to express their sincere thanks to the Director, Botanical Survey of In- 
dia for providing facilities and encouragement. The authors are also grateful to VJ. Nair, 
ex Joint Director, Botanical survey of India, Co-ordinator, AICOPTAX, Centre for Research 
on Grasses and Bamboos, Botanical Survey of India, Coimbatore (Tamilnadu), India, for 
providing Latin diagnosis. 
REFERENCES 

Tresster hi PVs Fnyrogenysanta Uassiitediton Ur bre wre amy: Diosconiae'rress, Fortianta; Uk 
Pearce, N.R.and PJ. Criss. 2002. The orchids of Bhutan. In: Flora of Bhutan 3(3):109. 


SINGH, K.P, S. PHUKAN, and P. Busarsarua. 2002. Orchidaceae. In: N.P. Singh and D.K. Singh, eds. Floristic 
diversity and conservation strategies of India 4:1735-1846. BSI, Calcutta. 


FOUR NOVELTIES AND A LECTOTYPIFICATION IN MATELEA 
(APOCYNACEAE: ASCLEPIADOIDEAE) FROM HISPANIOLA 


Alexander Krings 


Herbarium, Department of Plant Biology 
North Carolina State University 
Raleigh, North Carolina 27695-7612, U.S.A. 
Alexander_Krings@ncsu.edu 


ABSTRACT 


On- “going study of subtribe Gonolobinae (Apocynaceae: Asclepiadoideae) in the West Indies resulted in the dis- 
covery of four new species from Hispaniola: Matelea corynephora, M. phainops, M. rhynchocephala, and M. 
torulosa. Matelea pauciflora is cectoty pitied. 


RESUMEN 


me GescnDen © cuatro ae nuevas de Matelea de la Espanola como resultado del estudio de la subtribu 


a Asclepiadoideae) en las Indias Occidentales: Matelea corynephora, M. phainops, M. 
fea hocepkals: y M. torulosa. Se lectotipifica Matelea pauciflora. 


Gonolobus pauciflorus Spreng. (Apocynaceae: Asclepiadoideae) was described in 1824 

ased on a Bertero specimen from Hispaniola (P!). Schlechter (1899) recognized the spe- 
cies, noting that at the time, it still remained known only from the Bertero collection. 
Erik Ekman subsequently contributed numerous new collections that were referred to 
G. pauciflorus by Ignatz Urban of B (fide annotations) and later accepted as well by Alain 
H. Liogier (fide annotations), author of the most recent treatment of Asclepiadoideae for 
Hispaniola (Liogier 1994). Woodson (1941) transferred G. pauciflorus to Matelea Aubl. as 
one of 105 new combinations that resulted ina very broadly circumscribed genus thatis, 
in large part, still currently followed (see Stevens 2005). According to Woodson’s (1941) 
generic concept, Matelea is distinguished from Gonolobus Michx. in part by the absence 
of laminar dorsal anther appendages (Cd sensu Kunze 1995) in the former. A seemingly 
related species—Matelea linearipetala Alain—was described in 1988 from a single 1926 
Haitian specimen (Ekman H7136, S!) not seen by Urban. However, in the course of on- 
going systematic study of subtribe Gonolobinae in the West Indies, it has become clear 
that M. pauciflora (Spreng.) Woodson sensu Urban and Liogier contains four additional 
undescribed species. The four new species are described and illustrated below. For clar- 
ity, amplified descriptions, as well as illustrations, are also provided for M. linearipetala 
and M. pauciflora. The six species are nearly indisti e in the absence of flow- 
ers—each peaMine small, opposite, ovate-oblong leaves— but differ dramatically in vari- 
ous floral characteristics. A key follows. Corona terminology primarily follows Liede and 
Kunze (1993) and Kunze (1995), although it is recognized that additional work is needed 
to clarify family-wide homology (see Endress & Bruyns 2000; Livshultz 2003). 


1. Calyx colleters 2 per sinus; long eglandular sharp trichomes (ca. 0.2 mm) of the pedicels ret- 

rorse; corolla lobes ocellate (i.e, each bearing a single white spot on the adaxial surface near 

the apex); gynostegial corona (C(is)) columnar at the base and appearing as a ca. 0.6 mm tall 

stipe; stylar head lacking a terminal appendage M. phainops 
1. Calyx colleters 0-1 per sinus;long eglandular sharp trichomes of the pedicels predominantly, if 


SIDA 22(2): 941-953. 2006 


942 BRIT.ORG/SIDA 22(2) 


not exclusively, antrorse or spreading; corolla lobes ocellate or not; gynostegial corona (C(is)) 
not basally columnar nor appearing as a stipe; stylar head with or without a terminal appendage 
2. eee: corona (Clis)) with a broad, hardening, lobe-like, translucent yellow- en se- 
cretion in the interstaminal position (i.e, between the raised portions of the c a seg- 
ments at rise to meet the lower portion of the ee in the staminal position); ea 
nostegia na (C : unlobed; anther wings horizontally divergent; stylar head exhibiting 

5 ati en “ns age 
3. Gynostegial corona ci ) ca. 0.6-0.7 mm tall, staminal gynostegial corona (Cs) unlobed 
apically (i.e., descending smoothly in an arc from the apical ridge to the base); anther 
ae slightly longer than the corpusculum; terminal appendage of stylar head absent, 


+ 
— 


etime a short Scares Lene to 0.2 mm tall M. linearipetala 
| (C(is)) > 1 tall, inal gynostegial corona o slightly vertically 
“lobed apieally (.e., each ridge Le rising vertically immediately before descending to 


the base); anther wings slightly shorter than the corpusculum; terminal appendage of 
stylar head conic (volcano-shaped; broader at the base than at the apex),0.9-1.2 mm tall 
M. pauciflora 
2. Gynostegial corona (C(is)) lacking a hardening, lobe-like, translucent yellow-golden secre- 
tion in the interstaminal position; interstaminal gynostegial corona (Ci) lobed (or if not, then 
the stylar head with a knob-shaped terminal appendage); anther wings horizontally conver- 
gent at least at the apex or parallel; stylar head exhibiting a terminal appendage 
4. Calyx colleters absent; corolla lobe bases each with two round, distinct, lateral thicken- 
ings, basal lobe surfaces concave ("scooped-out’); interstaminal gynostegial corona (Ci) 
unlobed, staminal gynostegial corona (Cs) apically ungrooved, unlobed laterally, scallop- 
shaped in broad outline, margins entire or somewhat undulating; anther wings parallel; 
terminal appendage of stylar head capitate (e.g., reminiscent of a round door knob), to 1.1 
mm tall M. torulosa 
4.Calyx colleters 1 per sinus;corolla lobe bases lacking round lateral thickenings, lobes plane; 
interstaminal gynostegial corona (Ci) shallowly to broadly lobed, staminal gynostegial 
corona (Cs) apically grooved or laterally lobed, not scallop-shaped; anther wings conver- 
gent at least apically; terminal appendage of stylar head clavate or narrowly elongate- 
conic (obelisk-shaped), > 1.8 mm tall. 
5. Interstaminal gynostegial corona (Ci) a large lobe, staminal gynostegial corona (Cs) 
grooved apically, lacking two broad, lateral lobes; terminal appendage of stylar head 
clavate . corynephora 
5. Interstaminal gynostegial corona (Ci) a shallow lobe, staminal agynostegial corona (Cs) 
apically not grooved, broadly lobed laterally to appear 3-lobed; terminal appendage of 
stylar head narrowly elongate-conic (obelisk-shaped) M. rhynchocephala 


Matelea corynephora Krings, sp. nov. (Fig. 1). Type: HAITE Ile la Tortue, la Vallée, top of Morne Bar- 
ranca, ca. 300 m, 21 Mar 1928 (£1), E.L. Ekman H9740 (HOLOTYPE: S!). 


Species nova Matelea Aubl. diftert a Matel ill ilol i i lol 
eee h 


Slender, woody vine. Latex unknown. Stems ca. 1.8 mm diam., pubescent, glandular-capi- 
tate trichomes throughout, ca. 0.09 mm long, eglandular sharp trichomes mostly in 2 
lines, one on each side of the stem, retrorse, ca. 0.4 mm long. Leaves opposite, simple, blades 
oblong to linear-oblong, (7.2-)12-25 « (3.6-)5.2-8.6 mm, apex obtuse, apiculate, base 
broadly cuneate to rounded, margins entire, adaxial surface dark green, glossy on drying, 
glabrous or minutely pubescent along midvein with short glandular-capitate trichomes, 
abaxial surface pubescent along midvein and sometime secondary veins, trichomes 
mostly glandular-capitate, eglandular sharp trichomes few and scattered, colleters 2; 
petioles 3.8-7 = ca. 0.6 mm, glandular-capitate trichomes dense, ca. 0.1 mm long, 
eglandular sharp trichomes dense, antrorse, ca. 0.2 mm long, both trichome types found 
throughout, but the longer sharp trichomes tending to be slightly denser along the adaxial 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 943 


po 


Fic. 1. Motel } a A hah; 1 dotail nflaat-D <I re | tat H | 4 Y doy ee | } 1; C, gynostegial 
7 


corona—note the laterally unlobed Cs, and the rounded lobe a the i D, corpusculum and anther wings; E polinarium: caudicle 
shape appioumates Based on aD eae (S). aw = anther wings; Ci gy F ; 


P J) y A hte vA LULL ne Pr J 


petiolar ridge; stipular colleters 2, one on each side of petiole base, ca. 0.3 mm long. Inflo- 
rescence racemiform, peduncles 0.6-1.7 x ca. 1mm, glandular capitate trichomes absent, 
eglandular sharp trichomes 0.2-0.3 mm long, antrorse; pedicels 3-5 x ca. 0.6 mm, glan- 
dular-capitate trichomes dense, ca. 0.04 mm long, longer eglandular sharp trichomes 
dense, ca. 0.2 mm long, antrorse. Calyx 5-lobed, lobes oblong-ovate, ca. 0.9 x 0.6 mm, api- 
ces obtuse, margins entire, adaxial surface glabrous, abaxial surface very sparsely pubes- 
cent with eglandular trichomes scattered near center and along the margins: colleters 1 
per sinus. Corolla subrotate, bases campanulate, tube ca. 0.9-1.2  1.2-1.3 mm, 5-lobed, 
lobes imbricate in bud (not or only slightly dextrorse), green (fide collectoris), lanceolate- 
linear, 4.5-5.5 x 1.6-1.8 mm, apex obtuse, at least some ocellate (i.e., each lobe bearing a 
white eye adaxially at its apex), adaxial surface glabrous, abaxial surface glabrous or very 


944 BRIT.ORG/SIDA 22(2) 


sparsely pubescent with trichomes eglandular. Faucal annulus (corolline corona or Ca) 
indistinct. Gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), ca. 0.7 
mm tall, not appearing as a stipe, Cs rising to meet the bottom of the anther, rising seg- 
ment lacking two broad, lateral lobes, but grooved apically, Ci broadly lobed; laminar 
dorsal anther appendages (Cd) absent. Pollinarium: corpuscula dark reddish-brown, ca. 
0.15 mm long; pollinia borne horizontally or with the cavity slanting upwards at an angle, 
ovate, ca. 0.37 x 0.27 mm. Stylar head witha terminal appendage, appendage clavate, 1.9- 
2.2 mm long, 1.4-1.5 mm wide at the apex, generally broader at the apex than at the base, 
papillate throughout, a tuft of eglandular sharp trichomes present in the center of the 
apex, trichomes ca. 0.3 mm long. Follicles unknown. 

Distribution. —Apparently endemic to Haiti [known only from the typel. 

Etymology.—The specific epithet refers to the conspicuous, clavate, terminal stylar 
head appendage. 

Discussion.—Matelea corynephora appears closely related to M. rhynchocephala (Fig. 
5) and M. torulosa (Fig. 6)—both latter species also exhibiting a stylar head with a termi- 
nal appendage. All three can be easily distinguished by the shape of the gynostegial co- 
rona, as well as that of the appendage (see key above and Figs. 1, 5-6). All three lack the 
broad, translucent yellow-golden, hardening secretions in the interstaminal position as 
seen in M. linearipetala (Fig. 2) and M. pauciflora (Fig. 3). 


— linearipetala Alain, Phytologia 64:346. 1988. (Fig. 2). Type: HAITL on olig. limestone, ca. 

200 m, Massif des Matheux, Thomazeau, Morne a Cabrits, a vine, fr. “luteis,” 24 Oct 1926 (f1 & fr), ELL. 

Ekman H7136 (HOLOTYPE: S!) [n.v. Urban. 

Slender, woody vine. Latex unknown. Stems ca. 0.9 mm diam., pubescent in two lines, 
less densely so with age, glandular-capitate trichomes, ca. 0.08 mm long, eglandular sharp 
trichomes to 0.45 mm long, retrorse. Leaves opposite, simple, blades oblong, oblong-ovate, 
or ovate, 17-45 x 9-21 mm, apices obtuse to rounded, apiculate, bases rounded to trun- 
cate, margins entire, adaxial surface dull glabrous, except minutely pubescent along the 
midvein, glandular-capitate trichomes ca. 0.08 mm long, eglandular sharp trichomes 
absent, abaxial surface glabrous, except minutely pubescent along the veins, glandular- 
capitate trichomes ca. 0.08 mm long, eglandular sharp trichomes very sparse if present, 
ca. 0.2 mm long, colleters 2; petioles 4.4-17 x ca. 0.4 mm, densely pubescent throughout, 
glandular-capitate trichomes ca. 0.07 mm long, eglandular sharp trichomes very sparse, 
ca. 0.3 mm long, antrorse to antrorse-appressed; stipular colleters 2, one on each side of 
petiole base, ca. 0.3 mm long. Inflorescence racemiform, peduncles 0.6-2 x ca. 0.7 mm, 
essentially glabrous; pedicels 2.5-5 x ca. 0.7 mm, densely pubescent throughout, glandu- 
lar-capitate trichomes ca. 0.08 mm long, eglandular sharp trichomes absent or more or 
less uniform throughout, though sparsely so, ca. 0.18 mm long, predominantly antrorse 
or spreading. Calyx 5-lobed, lobes lanceolate to ovate-oblong, 1.7-2 x 0.8-1.2 mm, apices 
obtuse, pubescent, glandular-capitate trichomes ca. 0.08 mm long; colleters | per sinus. 
Corolla subrotate, bases subcampanulate, tube ca. 0.8 X 1 mm, 5-lobed, lobes imbricate 
in bud (not or only slightly dextrorse), green (fide collectoris), apparently not ocellate, 
linear lanceolate, 4.8-6.5 X 1.6-2 mm, apices obtuse, margins entire, adaxial surface gla- 
brous, reticulate at least on drying, abaxial surface g is, except tube pubescent, glan- 
dular-capitate trichomes ca. 0.08 mm long. Faucal annulus (corolline corona or Ca) in- 
distinct. Gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), Cs rising 
to meet the lower portion of the anther, rising segment a narrow ridge, triangular when 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 945 


ee 
co * si 


() 
0.09 mm| LAAN 


3.7 cm 


Alain-A hahit diastdA wR flawor rae | n I 4 L . c 
o ! be Bi 


Fic. Mate] pollinarium 
) 1 LOTT g5,E, f ! 
PH ial 


+ c Z Dp A al P vu749 Oo Uangc7/c\ ah +4 r 
I PE follicle 130 &N7UO/ (5). AW = anther wings; Ul gy g 
| Ha lah l c yet ee a] lini 

lobes; co rpuscuium,; Cs gy g rf polinium, 


viewed directly from the front, neither laterally nor vertically lobed, ca. 0.67 mm tall, Ci 
unlobed, exhibiting a broad, hardening, lobe-like, translucent yellow-golden, secretion; 
anther wings with divergent apices, only nearly touching at the base immediately sub- 
tending the corpusculum, vertical length immediately subtending corpusculum ca. 0.09 
mm long, horizontal length ca. 0.15 mm long; laminar dorsal anther appendages (Cd) 
absent. Pollinaria: corpuscula dark reddish-brown, ca. 0.15 mm long: pollinia borne hori- 
zontally, ovate, ca. 0.34 X 0.2 mm. Stylar head lacking a pronounced terminal append- 
age, sometimes exhibiting a slightly raised, circular or angular protuberance, ca. 0.2 mm 
tall. Follicle Gmmature?) 3.7 x 1.4. cm, 5-winged, wings interrupted and appearing as den- 
tate lobes, surface appearing glabrous, but minutely papillate throughout. Seeds unknown. 
Distribution.—Apparently endemic to Haiti. 


Additional specimens seen: HAITI: Massif des Cabos, Les Gonaives, Petite-Gouyne, dry limestone region, hill- 
sides, in thickets, vine, fl. green, ca. 150 m, 26 Sep 1927 (£1), E.L. Ekman H9067 (Cl, GH!, S!) [Referred to G. pauciflorus 


946 BRIT.ORG/SIDA 22(2 


= 


| 


indicates Ekman’s suspicion 0 gh without “Seems to come close 
to G. pauciflorus, but the umabell ae are cnoetly one-flowered and the sedunele very short.”} Ile la Gonave, Pte-a- 
Raquettes, hillsides, f1. oe sisaeaes (ED, ELL. jail H8809 (US|, S!) [Referred to G. pauciflorus by Urban and 
s handwritten note in pencil on the S sheet adds to the locality 


by Urban and M. pauciflora by L cas a annotations. A | handwritten note in Na by Ekman on the S sheet 
fi 


M. pauciflora by Liogie 
information: “I have not analyzed the flowers but the ge pas seem to aan any well wat nls IS 
pauciflorus]. Better plant. Coll. before!” However,“ pauciflorus” 


above the epithet, suggesting perhaps a later second-guess by al . 
Matelea pauciflora (Spreng.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. (Fig. 3). 
BASIONYM: Gonolobus pauciflorus Spreng., Systema Vegetabilium, editio decima sexta 1:846. 1824. TYPE: 
HISPANIOLA: ‘St. Dominique, Bertero s.n. (LECTOTYPE, here designated: P!). 
Slender, woody vine. Latex unknown. Stems ca. 1.7 mm diam., pubescent throughout or 
in two lines, glandular-capitate trichomes ca. 0.07 mm long, eglandular sharp trichomes 
ca. 0.3mm long, retrorse. Leaves opposite, simple, blades oblong or ovate, 8.5-56.5 x 5.3- 
27.8 mm, apices obtuse, bases rounded to truncate, margins entire, adaxial surface dull, 
glabrous except minutely pubescent along midvein, glandular-capitate trichomes ca. 0.05 
mm long, eglandular sharp trichomes sparsely scattered, ca. 0.18 mm long, abaxial sur- 
face glabrous except pubescent along the veins, glandular-capitate trichomes ca. 0.08 
mm long, eglandular sharp trichomes sparsely scattered, ca. 0.4 mm long, colleters 2; 
petioles 3.6-22.2 x ca. 0.4 mm, densely pubescent throughout, glandular-capitate tri- 
chomes ca. 0.1 mm long, eglandular sharp trichomes scattered throughout, ca. 0.23 mm 
long, predominantly antrorse or spreading; stipular colleters 2, one on each side of peti- 
ole base, ca. 0.2 mm long. Inflorescence racemiform, peduncles to 5 x ca. 0.4 mm, gla- 
brous or pubescent, glandular-capitate trichomes ca. 0.08 mm long, eglandular sharp 
trichomes sparse, ca. 0.36 mm long, antrorse or spreading; pedicels to 9 x ca. 0.4 mm, 
densely pubescent, glandular-capitate trichomes ca. 0.05 mm long, eglandular sharp tri- 
chomes ca. 0.3 mm long, predominantly antrorse or spreading. Calyx 5-lobed, lobes lan- 
ceolate to lance-ovate, 1.3-2.5 x 0.4-1 mm, apices obtuse, bases pubescent, glandular- 
capitate trichomes ca. 0.05 mm long, margins with eglandular sharp trichomes ca. 0.1 
mm long, otherwise glabrous; colleters 1 per sinus. Corolla subrotate, bases 
subcampanulate, tube ca. 0.9 X 1.2 mm, 5-lobed, lobes imbricate in bud (not dextrorse), 
green (fide collectoris), apparently not ocellate, linear-lanceolate, 4.7-7.5 x 2-2.8 mm, 
apices obtuse, margins entire, adaxial surface glabrous, abaxial surface glabrous, except 
tube pubescent, glandular-capitate trichomes ca. 0.05 mm long. Faucal ; lus (corolline 
corona or Ca) indistinct. Gynostegial corona of fused staminal (Cs) and interstaminal 
(Ci) parts, Cs rising to meet the lower portion of the anther, rising segment swollen at the 
base, slightly vertically lobed along the apical ridge, ca. 1.25 mm tall, Ci unlobed, exhib- 
iting a broad, hardening, lobe-like, translucent yellow-golden, secretion; anther wings 
with divergent apices, only nearly touching at the base immediately subtending the 
corpusculum, vertical length immediately subtending corpusculum ca. 0.08 mm long, 
horizontal length ca. 0.18 mm; laminar dorsal anther appendages (Cd) lacking. 
Pollinarium: corpuscula dark reddish-brown, 0.18-0.2 mm long; pollinia borne horizon- 
tally orata slightly upwards angle, oblong, ca. 0.42 x 0.24 mm. Stylar head witha termi- 
nal appendage, appendage volcano-shaped, ca. 0.9-1.2 mm tall. Follicle unknown. 
Distribution —Haiti and Dominican Republic. 
Discussion.—Although bearing a conic terminal appendage of the stylar head, M. 
pauciflora (Fig. 3) appears more closely related to M. linearipetala (Fig. 2) than the other 
three terminally-appendaged species—M. corynephora (Fig. l), M. rhynchocephala (Fig. 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 947 


ee 
E Po 0:56 mini 


Fic. 3. Matelea pauciflora (Spreng.) Woodson: A, habit and leaf detail; B, flower with conic terminal appendage of stylar head; C, 
gynostegial corona—note slight vertical lobe of each Cs ridge kartowiea), “ corpusella and antic wings; E, mpallinanis caudicle 
shape aphfoxtmates Based on Exmiate Hol 71 (K). aw = anther wings; Ci gy g ; ; 


| Wi + + inal d £ ety kh | 


COPPUSCUIUTTT, & Yy y rv PUL yd Pt y y 


5), and M. torulosa (Fig. 6)—based on the shape of the gynostegial corona, the presence of 
broad, hardening, lobe-like, translucent yellow-golden, secretion in the interstaminal 
position, and the shape of the anther wings. Matelea linearipetala lacks a terminal ap- 
pendage of the stylar head and can be additionally distinguished from M.pauciflora by 
the height and shape of the aan corona (Fig. 2). 


Additional specimens seen: . d@Artibonite: in a little valley not far from Mirabalais, shrubby 
Laubwald, 26 Oct 1924 (£D, e E re oe a (GHI, IJ, NY!, US) [The sheet at US additionally includes the text: 
“on roadside, rare, 150 m.” Annotations referring the specimens here appear by Urban, Woodson, and Liogier. 
Ekman slips in pencil are lacking.|; Massif du Nord, Hinche, Morne Juan-Gomez, oligocene limestone, in thickets, 
ca. 700 m, 23 May 1926 (fl), E.L. Ekman Hel (Kl, = i vel ae the collection only to Poicillopsis, Ekman’s 

handwritten note in pencil on the S sheet i 1 the taxon. This specimen is the second 


948 BRIT.ORG/SIDA 22(2) 


earliest of his collections in the M. pauciflora “complex.” He is correct in noting “same as n. 2266.”]. DOMINICAN 
REPUBLIC. Prov. Azua: Cordillera cont ral, 5 kmalS ms eblo de Padre Las Casas, en una lomita situada en el 
lado E de la carretera hacia Las Yay iphus, Agave y Bursera, 18° 41'N, 70° 55'W elev. 680-700 
m, 2 Oct 1987 ({)), R. Garcia y J. pee 2531 oe ae iow to Matelea prior to this study], Bani, vine 

flowers greenish, in thickets, 24 Sep 1966 (fD), Bro. B.A. Lavastre 2290 (NY!) [Referred here by Liogier fide annota- 


tion]. 


Matelea phainops Krings, sp. nov. (Fig. 4). Tyre: DOMINICAN REPUBLIC: vine, up to 1.5 m high, flow- 
rs yellowish green, in thickets, about 2 mi W of Oviedo, alt. about sea level, on limestone, 3 Nov 1989 (£1), 
AH. Liogier 16617 (HOLOTYPE: GH} ISOTYPES: NY!, USF). 


Species nova Matelea Aubl. differt a Matelea antillanarum forma coronae et gynostegi (v. Fig. 4). 


Slender, woody vine. Latex unknown. Stems ca. 2.12 mm diam., pubescent, glandular- 
capitate trichomes throughout, ca. 0.12 mm long, eglandular sharp trichomes mostly in 
2 lines, one on each side of the stem, retrorse, ca. 0.36 mm long. Leaves opposite, simple, 
blades oblong or ovate, (8-)12-31 x (-4)6-14 mm, apices obtuse, apiculate, bases broadly 
cuneate to rounded or subtruncate, margins entire, adaxial surface dull to somewhat 
glossy on drying, glandular-capitate trichomes throughout, but most dense along the 
veins, ca. 0.13 mm long, eglandular sharp trichomes scattered on midvein or absent, to 
0.4mm long, abaxial surface glabrous, except for short glandular-capitate trichomes along 
the vines, ca. 0.05 mm long, colleters 2-3; petioles to 10 x ca. 0.6 mm, glandular-capitate 
trichomes dense, ca. 0.1 mm long, eglandular sharp trichomes occassional or scattered, 
antrorse, ca. 0.22 mm long; stipular colleters 2, one on each side of petiole base, ca. 0.23 
mm long. Inflorescence racemiform, peduncles to ca. 2.5 X 0.6 mm, glandular-capitate 
trichomes dense throughout, ca. 0.08 mm long, eglandular sharp trichomes scattered, 
spreading or antrorse, 0.17-0.2 mm long; pedicels 4-7 x ca. 0.36 mm, short trichomes 
both capitate and sharp, dense, ca. 0.09 mm long, longer eglandular sharp trichomes, if 
present, scattered and retrorse, ca. 0.2 mm long. Calyx 5-lobed, lobes ovate, 0.85-1.3 x 
0.6-0.9 mm, apices obtuse, margins entire, glabrous, adaxial surface glabrous, abaxial 
surface minutely glandular pubescent; colleters 2 per sinus. Corolla subrotate, bases 
subcampanulate, tube ca. 0.4 X 0.5 mm, 5-lobed, lobes imbricate in bud (not or only 
slightly dextrorse), oblong-linear, 2.5-2.7 x 1.2-1.4 mm (width at base), apex obtuse, yel- 
lowish green (fide Page) each lobe bearing a white eye at its apex, adaxial surface gla- 
brous, abaxial surf ,except tube pubescent, glandular trichomes ca. 0.09 mm. 
Faucal (corolline corona or Ca) indistinct. Gynostegial corona of fused staminal 
(Cs) and interstaminal parts (Ci), basally fused and appearing as a stipe, ca. 0.6 mm tall, 
upper Cs undulating, rising to meet the bottom of the anther in the staminal position, Ci 
unlobed; laminar dorsal anther appendages (Cd) absent. Pollinarium: corpuscula dark 
reddish-brown, ca. 0.13 mm long; pollinia borne horizontally, ovate, ca. 0.4 x 0.2 mm. 
Stylar head lacking a terminal appendage. Follicles unknown. 

Distribution.—Apparently endemic to the Dominican Republic [known only from 
the typel. 

Etymology—This vine is named in honor of my wife, the ‘Bright eyes’ in my life. The 
epithet also reflects the ocellate corolla lobes exhibited by this species. 

Discussion.—Matelea phainops appears related to the Cuban Matelea oblongata, M. 
acuminata, and M. nipensis (Urb.) Woodson and, perhaps, Jamaican Jacaima costata (Urb.) 
Rendle. It is one of several mostly West Indian taxa exhibiting ocellate corolla lobes. Con- 
tinental representatives likely include M. pusilliflora WO. Williams, M. ocellatus WD. 
Stevens, and the recently described M. emmartinezii WD. Stevens. In addition to the afore- 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 949 


aw 


10.18 mm 


eee Ne 


AK 
0.54 mm 


R flowe t Ilat Ila lob 1 ith like hace: D 
TIARG Vase, 


COTPUSCUIUTTT, Yy y ' Hae, p PUHTIUTTT, 


ML ioled aBainane KHAHeOR: ahitand eat ee p 5 
corpuscula and anther wings; E, pollinarium, caudicle shape approximate. Based on Liogier 16617 (GH). aw = anther wings; Ci = 
. _ 1 + or 1 1 ‘a + MH | + am Il lini 


mentioned, small flowered species, larger flowered taxa likely belonging to the complex 
in the West Indies include M. bayatensis (Urb.) Woodson, M. correllii Spellman, and M. 
tigrina (Griseb.) Woodson. The latter may not be specifically distinct (see Krings 2005). 
The morphology of the gynostegial corona is variable in the group—from indistinct to 
well-developed—and additional research is necessary to clarify specific relationships. 

Matelea rhynchocephala Krings, sp. nov. (Fig. 5). Tyee: DOMINICAN REPUBLIC. PROV. SANTIAGO: 


Valle del Cibao, Santiago, Hato del Yaque, in thickets, f1. green, fruiting, 15 Feb 1930 (f1 & fr), EL. Erman 
H14296 (HOLOTYPE: S!) [n.v. Urban; referred to M. pauciflora by Liogier fide annotation]. 


Species nova Matelea Aubl. differt a Matelea antillanarum corona parvilobata interstaminer, lobata staminer, 
a o a! } 1 


gyt 


950 BRIT.ORG/SIDA 22(2) 


2.9 mm 


46cm 


Fic. 5 Motel, hy a pt lo Ue g A,| hi lin-fd aT B, flower; C gy + gi 1 + ee ad chal Ink £ 
Ci; D, corpuscula and anther wings; E, pollinarium, caudicle shape approximate. Based on Ekman H14296 (S).aw = sanmnier einige: c= 
: . n oot | Ila Inh | C | 

rolla lobes; p : gynosteg po = pollinium ;ta=termi 


nal appendage of stylar head. 


Slender, woody vine. Latex unknown. Stems ca. 1.2 mm diam., densely pubescent through- 
out, though 2 lines somewhat evident on older stems, glandular-capitate trichomes ca. 
0.06 mm long, eglandular sharp trichomes ca. 0.4 mm long, retrorse or spreading. Leaves 
opposite, simple, blades oblong or ovate, 19-48 x 7-20 mm, apices obtuse, apiculate, bases 
rounded to truncate, margins entire, adaxial surface dull on drying, not glossy, glabrous 
or pubescent along the midvein, glandular-capitate trichomes ca. 0.1mm long, eglandular 
sharp trichomes absent or essentially so, abaxial surface glabrous, pubescent along the 
veins, glandular-capitate trichomes ca. 0.1 mm long, eglandular sharp trichomes to 0.2 
mm long, colleters 2; petioles to 14 x ca. 0.45 mm, densely pubescent, glandular-capitate 
trichomes ca. 0.07 mm long, eglandular sharp trichomes ca. 0.3 mm long, throughout, 
antrorse; stipular colleters 2, one on each side of petiole base, ca. 0.2 mm long. Inflores- 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 951 


cence racemiform, peduncles ca. 0.8 x 0.5mm, pubescence as that of stems; pedicels 1.7- 
18 X ca. 0.44 mm, densely pubescent, glandular-capitate trichomes ca. 0.09 mm long, 
eglandular sharp trichomes ca. 0.2 mm long, antrorse or spreading. Calyx 5-lobed, lobes 
lanceolate, 1.5-1.6 x 0.5-0.6 mm, glandular-capitate trichomes mostly at the base, ca. 0.07 
mm long, glandular sharp trichomes mostly near the middle, ca. 0.2 mm long; colleters 1 
per sinus. Corolla subrotate, bases subcampanulate, tube ca. 1 X 1 mm, 5-lobed, lobes im- 
bricate in bud (not or only slightly dextrorse), green (fide collectoris), linear-lanceolate, 
4.9-5.9 x L.1-1.4 mm, apex obtuse, margins white, adaxial surface glabrous, abaxial sur- 
ly pubescent, glandular-capitate trichomes ca. 0.07 
mm long. Faucal annulus (corolline corona or Ca) indistinct. Gynostegial corona of fused 
staminal (Cs) and interstaminal (Ci) parts, Cs rising to meet the lower portion of the an- 
ther, rising segment with a single, broad lobe on either side, ca. 0.8-0.9 mm tall, Ci shal- 
lowly lobed; anther wings triangular in frontal view, nearly touching; laminar dorsal 
anther appendages (Cd) absent. Pollinarium: corpuscula dark reddish-brown, ca. 0.3mm 
long, pollinia borne horizontally or with the cavity slanting upwards at an angle, ovate, 
ca. 0.4 X 0.25 mm. Stylar head with a terminal appendage, appendage narrowly elon- 
gate-conical (obelisk-shaped), 2.8-3.3 X 0.6-0.8 mm, papillate, apex obtuse, with a tuft 
of eglandular trichomes or glabrous. Follicles ovate, 5.3-1.9 cm, 5-winged, wings inter- 
rupted and appearing as dentate lobes, glabrous. Seeds ovate, ca. 4.9 x 3.4 mm, glabrous, 
margins smooth, neither warty, nor dentate. 

Distribution.—Apparently endemic to the Dominican Republic. 

Etymology.—The specific epithet refers to the obelisk-shaped or snout-like terminal 
appendage of the stylar head. 

Discussion.—Matelea rhynchocephala (Fig. 5) appears most closely related to M. 
corynephora (Fig. 1) of Haiti. Notable differences between the two are the shape of the 
terminal appendage of the stylar head (obelisk-shaped in M. rhynchocephala vs. clavate 
in M. corynephora) and the shape of the gynostegial corona (Figs. 1 & 5). 


face glabrous in age. except tub 
oO o~™ C 


Additional specimen seen: DOMINICAN REPUBLIC: El Rubio, 22 Nov 1940 (£1), M. Canela s.n. (NY!) [Referred to 
M. pauciflora by Liogier fide annotation]. 


Matelea torulosa Krings, sp. nov. (Fig. 6). Type: DOMINICAN REPUBLIC. Distr. NACIONAL: Los 3 Ojos 


Agua, near Santo Domingo, 31 Oct 1959 ({D, E. Marcano [],J. Jiménez] 4096 (HOLOTYPE: US!) [referred to 
M. pauciflora by Liogier fide annotation]. 


Species nova Matelea Aubl. differt a Matel ill ( lo} ; : TAG 


gynostegio toruloso. 


Slender, woody vine. Latex unknown. Stems ca. 1.2 mm diam., densely pubescent in two 
lines, glandular-capitate trichomes ca. 0.05 mm long, eglandular sharp trichomes ca. 0.4 
mm long, retrorse. Leaves opposite, simple, blades oblong or ovate, 9-28 X 3-8.8 mm, api- 
ces obtuse, apiculate, bases rounded or broadly cuneate, margins entire, adaxial surface 
dull, glabrous except pubescent along the midvein, glandular-capitate trichomes ca. 0.07 
mm long, eglandular sharp trichomes sparse, to 0.18 mm long, abaxial surface pubescent 
primarily along the veins, glandular-capitate trichomes ca. 0.1mm long, eglandular sharp 
trichomes to 0.4 mm long, colleters 2-3; petioles 2-6 X ca. 0.4 mm, densely pubescent 
throughout, glandular-capitate trichomes ca. 0.05 mm long, eglandular sharp trichomes 
ca. 0.33 mm long, antrorse to antrorse-appressed, not spreading or rarely so; stipular 
colleters 2, one on each side of petiole base, ca. 0.22 mm long. Inflorescence racemiform, 
peduncles to 4.9 X ca. 0.7 mm, densely pubescent throughout, glandular-capitate tri- 


952 BRIT.ORG/SIDA 22(2) 


0.53 mm 


AK 


Fic. 6. Matelea torulosa Krings: A, habit; B, flower t 
forming a distinct pit (arrowed); C, stylar head Jage; D, ial 
diel hk 7 to R d AA 


, concave (“scooped-out”) and 
eer ings; E, pollinarium, 
ety I 


rr J Jt 
ay . 


J J 
TANO dic +h 1 

7U70 (U>),aW = anther win gs; 
Haloh ! c Fee, eee tanial Wee: dane oh aa 
corolla lobes , CO = cOrpusculum,; © gy g rf pollinium, t PI g y head. 


chomes ca. 0.07 mm long, eglandular sharp trichomes ca. 0.2 mm long, predominantly 
antrorse, some retrorse, sparser than glandular-capitate trichomes; pedicels 4.5-6.1 x 0.4 
mm, densely pubescent, glandular-capitate trichomes ca. 0.07 mm long, eglandular sharp 
trichomes ca. 0.18 mm long, predominantly antrorse or antrorse appressed. Calyx 5-lobed, 
lobes lanceolate or lance-ovate, ca. 1.8 x 0.8 mm, apices obtuse, margins entire, scattered 
pubescent, glandular-capitate trichomes to 0.07 mm long, eglandular sharp trichomes 
to 0.2 mm long; colleters lacking. Corolla subrotate, bases subcampanulate, tube 0.9 x | 
mm, 5-lobed, lobes imbricate in bud (not dextrorse), linear-lanceolate, 4.5-5 x 1.3-16mm, 
apices obtuse, margins entire, marginally thickened at the base on each side, scooped- 
out and forming a distinct pit near the base, otherwise plane, adaxial surface papillate, 
abaxial surface glabrous or witha few, eglandular sharp trichomes ca. 0.2 mm long. Faucal 
annulus (corolline corona or Ca) indistinct. Gynostegial corona of fused staminal (Cs) 
and interstaminal (Ci) parts, Cs rising to meet the lower portion of the anther, rising seg- 


KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 953 


ment scallop-shaped in outline, ca. 0.5 mm tall, not lobed on either side, yet exhibiting a 
smaller segment immediately in front, margins somewhat undulating to entire, Ci 
unlobed; anther wings parallel, nearly touching, well-developed toward the center of the 
stylar head, ca. 0.4-0.5 mm long, the vertical segment immediately subtending the 
corpusculum ca. 0.2 mm long, slightly longer than the corpusculum or as long: laminar 
dorsal anther appendages (Cd) absent. Pollinarium: corpuscula dark reddish-brown, ca. 
1.8-1.9 mm long; pollinia borne horizontally, ovate, ca. 0.37 X 0.25 mm. Stylar head with 
a terminal appendage, appendage capitate (“door knob-shaped”), ca. 1-L1 mm tall, apex 
ca. | mm broad. Follicles unknown. 

Distribution.—Apparently endemic to the Dominican Republic [known only from 
the typel. 

Etymology—The epithet refers to the capitate, door knob-like terminal appendage 
of the stylar head. 

Discussion.—The appendage is among the shortest of the four species bearing them 
in Hispaniola. Only Matelea corynephora (Fig. )) exhibits an appendage similarly broader 
at the apex than at the base, but the appendage is much longer than in M. torulosa and 
clavate. The two species also differ in a number of additional floral characteristics, such 
as the morphology of the gynostegial corona. The other appendaged species—M. 
pauciflora (Fig. 3) and M. rhynchocephala (Fig. 5)—exhibit terminal appendages that are 
narrower at the apex than at the base. 


ACKNOWLEDGMENTS 


[thank the curators and staff of the following herbaria for searching, or providing access 
or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUE.C, CGE, COLO, 
CR, DUKE, E, F FI, FLAS, FR, FIG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, 
K, LD, M, MICH, MIN, MO, MSU, NEU, NSW, NY, O, OXE P, PH, RSA, S, U, UC, UPS, US, USE 
TUR, WILLI, WU, Z. l also thank Roxana Yockteng for her kind assistance during, and 
subsequent to, a visit to P. The North Carolina State University Libraries Inter-Library 
Loan Service was instrumental in obtaining necessary titles not held in the collections. 
Alain Liogier and an anonymous reviewer provided thoughtful reviews of a previous 
version of the manuscript. 


REFERENCES 


Enpress, M.E. and P.V. Bruyns. 2000. A revised classification of the Apocynaceae, s.|. Bot. Rev.66:1-56. 

Krincs, A. 2005. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - 
Asclepiadoideae - Gonolobinae) in the Greater Antilles and Bahamas. Sida 21:1525-1533. 

Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211-238. 

Liebe, S. and H. Kunze. 1993. A descriptive system for corona analysis in Asclepiadaceae and 
Periplocaceae. Pl. Syst. Evol. 185:275-284. 

Liocier, H.A. 1994. La flora de la Espanola, VI. Publ. Univ. Central Este. 70:1-517. 

LivsHuttz, T. 2003. Systematics of Dischidia (Apocynaceae, Asclepiadoideae). Ph.D. dissertation, Cor- 
nell University. 

SCHLECHTER, R. 1899. Asclepiadaceae. In: Symbolae Antillanae, |. Urban,ed.Gebriider Borntraeger, Berlin 
Pp. 236-290. 

Stevens, W.D. 2005. New and interesting milkweeds (Apocynaceae, Asclepiadoideae). Novon 15: 
602-619, 

Woooson, R.E.1941.The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28:193-244. 


954 BRIT.ORG/SIDA 22(2) 


Book REVIEW 


ANTHONY Huxtry. 2005. Green Inheritance: The WWF Book of Plants.SBN 0-520-24359- 
5, pbk.). The University of California Press, Berkeley, CA 94704, U.S.A. (Orders: Cali- 
fornia Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. 
Tel: 609-883-1759; Fax 609-883-7413, www.ucpress.edu). $29.95, 192 pp., 234 color, 
OE LO fe 


Why is environmental conservation important? If you have ever asked this question, this book will provide your 
answer. Focusing on the “green mantle” of plant life that surrounds us, ie ne lela ae reader how conser- 


vation of the environment is integral to human survival. Color pl inc 


uding botani- 


cal illustrations from the collection at Kew Gardens, complement os clear writir 


] f 
ig style of the late Mr | Huxley, a 
prolific writer and an editor at Amateur Gardening. In this book, Mr. Huxley reminds us that “Without plants, 


animal lif Id ist.” From the very beginning, plants have acted as the “life support systems” of the planet, 
providing human and animal life with protection, nourishment, and even the air we panes It was the emer- 


gence of green plants that changed the poisonous primeval atmosphere into today’s | ygen-bearing 
atmosphere. In Chapter 2, “Guardians of the Environment”, the role of plants in protecting the air we breathe, 


preventing soil erosion, protecting us from harmful radiation, fixing nitrogen in the soil, stabilizing the climate, 


and producing rain is discussed. The role of cultivated plants in providing us with food and spices is celebrated, 
as well as the problems resulting from modern agricultural techniques. The 5,000-year history of “green medi- 
cine” is astanee from traditional pera! cures to modern drug aIScONeEy ao orts ane the resurgence of herbal 
| 1.Inachapter titled “Plants 
and Seeley" the sp aia of pants in our be: day lives and in the lives of other cultures is discussed. Mr. 


medicine f medicinal f 


t 


Huxley pter to the roles of in situ conservation, such as land reserves, and ex situ conserva- 
tion, such as modern bio- i pinnae and gene hanks, in protecting our resources and f feeding a nungty P planet. 


The author reminds us throughout the book that * tho ight- 


less impatience and greed, failing to appreciate the value of what we ae The last ciapeer is appropriately 
titled “Saving the Plants that Save Us” and summarizes the environmental problems that are depleting plant life 

on our planet, as well as solutions. The primary approaches to saving our “green inheritance” include halting 
environmental degradation, rehabilitating land, and conserving the pristine natural and semi-natural vegeta- 
tion that still remain. Mr. Huxley does not sugar-coat the issue, he acknowledges the difficulty of getting nations 
to work together, as well as the role that economic disparity plays in the battle. But, he paints a beautiful picture 
of the reward, “Besides ¢ thus providing for our io nee plants can also pone solace for the mind. They 


are endlessly fa Ascin AtiNg, we asia g word, and the ey are diversely beau- 
tiful too.”—Marissa Oppel, MS, C dR *h Assistant, Botanical Research Institute of Texas, Fort Worth, 


TX, 76102-4060, U.S.A. 


SIDA 22(2): 954. 2006 


NEW COMBINATIONS IN THE GENUS CYMOPTERUS (APIACEAE) 
OP THE SOUTHWESTERN UNITED STATES 


Ronald L. Hartman 


Rocky Mountain Herbarium 
Department of Botany 
University of Wyoming 

. University Ave. 

Laramie, Wyoming 82071-3165, U.S.A. 

Rhartman@uwyo.edu 


— 


ABSTRACT 


Preparation ol the pmiateas for Bolack San Juan Basin Flora has necessitated the following new combinations: 
C. davidsonii, C. glomeratus var. fendler i, and C. ses ili lorus 


eA 1 


RESUMEN 


La preparacion de las cane ee la Flora de Bolack San Juan Basin | do] combinaciones 


nuevas: Cymopterus breviradiatus, C. davidsonii, C. glomeratus var. fendleri, y C. eciils Orus. 


In revising Cymopterus Raf. (Apiaceae) of the greater Four Corner’s region for the San 
Juan Flora (Heil et al.), it has become necessary to make the following nomenclatural 
innovations. The aforementioned treatment represents a continued reliance on charac- 
ters of the fruit and of vegetative morphology (Mathias & Constance 1944-45; Cronquist 
et al. 1997). Recently, several phylogenetic analyses of nuclear (ITS) and plastid (rpsl6 
intron) sequences (Downie et al. 2002; Sun @ Downie 2004; Sun et al. 2004) have defined 
a group, likely monophyletic, of western North American apioids. Genera include Aletes 
J.M. Coult. & Rose, Cymopterus [here circumscribed to include Oreoxis Raf., 
Pseudocymopterus J.M. Coult. & Rose, and Pteryxia (Nutt. ex Torr. & A. Gray) J.M. Coult. 
& Rose], HarbouriaJ.M. Coult. & Rose, Lomatium Raf., Musineon Raf., Neoparrya Mathias, 
Oreogenia S. Wats., Oreonana Jeps., Podistera S. Wats.,and Shoshonea Evert & Constance. 
Thus far the molecular trees are poorly resolved with the basal branches containing sev- 
eral to many polytomies, thus obscuring relationships among terminal groups. A num- 
ber of terminal clusters make sense to one steeped in western umbels while others seem- 
ingly defy logic. It is hoped that meet pequcuce data will provide sufficient insight 
into the evolution of the western ay at a stable classification can be established. 


Cymopterus breviradiatus (W.L. Theob. &@ CC. Tseng) R.L. Hartm., comb. @ stat. nov. 
BASIONYM: Aletes macdougaliiJ.M. Coult. & Rose subsp. breviradiatus WL. Theob. & CC. Tseng, Brittonia 
16:306, fig. 6.1964. 


The genus Aletes is defined by its laterally Hoses (versus terete to dorsally flattened) 
fruit. This lateral compression is prominent in the type species, A. acaulis (Torr.)J.M. Coult. 
& Rose and its presumed closest relatively, A. humilis J.M. Coult. @ Rose. None of the 
three taxa of Aletes for which nomenclatural innovations are here made have fruit that 
are flattened laterally. Although molecular studies may eventually show that this is a 
fickle character, it is one that has been relied upon traditionally. 

Based on gestalt and technical features of the fruits and leaves, Cymopterus breviradiatus 
is extremely similar to Cymopterus sessiliflorus (Theobald et al. 1964) despite the fact 


SIDA 22(2): 955-957. 2006 


956 BRIT.ORG/SIDA 22(2) 


that the two taxa do not group near one another based on molecular data (Fig. 2, Sun & 
Downie 2004). There is ample justification for removing subsp. breviradiatus from sub- 
jugation to Cymopterus macdougalii J.M. Coult. & Rose) Tidestrom [Oreoxis m.].M. Coult. 
& Rose, 1913; Aletes m. J.M. Coult. & Rose, 1935]. The former has fruit with corky-thick- 
ened wings whereas C. macdougalii has weakly developed, thin wings. Cymopterus bechii 
S.L. Welsh & Goodrich appears virtually identical to C. macdougalii morphologically. 
This is reinforced thanks to the number of collections of each that have been amassed in 
recent years. Consequently the former is treated as a synonym of C. macdougalii. Fur- 
thermore, as sequence data indicate (Figure 2, Sun @ Downie 2004), C. macdougalii, C. 
(Pteryxia) davidsonii, and C. beckii group with C. lemmonii (Coult. & Rose) R.D. Dorn 
[Pseudocymopterus montanus J.M. Coult. & Rose]. C. davidsonii strongly resembles C. 
macdougalii morphologically and they may represent sister taxa. 


eS 


i davidsonii (J.M. Coult.& Rose) R.L. Hartm., comb. nov. Basiony: Aletes? davidsonii 

M. Coult. & Rose, Contr. U.S, Natl. Herb. 7:107. 1900. Peuaeconun we davidsonii J.M. Coult. & Rose) 

Mathias Ann. Missouri Bot. Gard. 17:282, 316. 1930. Pteryxia davidsonii .M. Coult. @ Rose) Mathias & 
Constance, Bull. Torrey Bot. Club 69:248. 1942. 


This species was most recently treated in Pteryxia (Mathias & Constance 1944-45) and 
tentatively attributed to Pseudocymopterus (Cronquist et al. 1997). 
The reader is referred to comments provided under Cymopterus breviradiatus. 


Cymopterus glomerattis (Nutt. ) DC. var. fendleri (A. Gray) R.L. Hartm., comb. & stat. nov. 


BASIONYM: Cy terus i A. Gray, Mem. Amer. Acad.,n 56.18 
Specific epithet: C sitentars glomeratus (Nutt. ) DC,, Prodr. 4: nen 1830. BASIONYM: Thapsia glomerata Nutt., 
Gen. N. Amer. Pl. 1:184. 1818, for which C s Raf., Herb. Raf. 2:40. 1833, is illegimate and 


— 


surperfluous (nom. nov. for Selinum eee Push FL Annee Sept. 2:732, ne non Cav, 1799; Internationa 

Plant Names Index 2005). 
Cymopterus fendleri long has been treated as distinct from the related C. glomeratus (C. 
acaulis, Mathias & Constance 1944-45) or as a variety of it (Cronquist 1997 et al; Goo- 
drich 2003). A recent study (Sun et al. 2005) using principal component analyses failed 
to discriminate among the five varieties recognized by Goodrich (2003). Despite this lack 
of resolution, the above combination is made for botanists, including myself, wishing to 
recognize var. fendleri. 


Cymopterus sessiliflorus (W.L. Theob. & C.C. Tseng) R.L. Hartm., comb. nov. BasionyM: Aletes 
sessiliflorus WL. Theob. & C.C. Tseng, Brittonia 16:309. 1964. 


The reader is referred to comments provided under Cymopterus breviradiatus. 


ACKNOWLEDGMENTS 


I appreciate the comments of an anonymous reviewer. 


REFERENCES 


Cronauist, A., N.H. Houmaren, and P.K. Houmaren. 1997. Intermountain flora: Vascular plants of the Inter- 
mountain West, U.S.A. Vol. 3A: Subclass Rosidae (except Fabales). New York Bot. Garden, Bronx. 
P. 372. 

Downie, S.R., R.L. HARTMAN, F.-J. Sun, and D.S. Katz-Downie. 2002. Polyphyly of the spring-parsleys 
(Cymopterus): molecular and morphological evidence suggests complex relationships among 
the perennial endemic genera of western North American Apiaceae. Canad. J. Bot. 80:1 295-1324. 


HARTMAN, NEW COMBINATIONS IN CYMOPTERUS 957 


GoooricH, S. 2003. In: S.L. Welsh, N.D. Atwood, S. Goodrich, and L.C. Higgins, eds. A Utah flora, ed. 3. 
Brigham Young Univ., Provo, Utah. P.654. 

Heit, K. et al. Bolack San Juan Basin flora. Missouri Botanical Garden Press, St. Louis, Missouri.In prep. 

INTERNATIONAL PLANT Name INoex, THE. 2005. (Attp://www.ipni.org) 

Maruias, M.E.and L. Constance. 1944-45. Umbelliferae. N.Amer.Fl.28B:187.New York Bot.Garden, Bronx 

Sun F-J. and S.R. Downie. 2004. A molecular systematic investigation of Cymopterus and its allies 
(Apiaceae) based on phylogenetic analyses of nuclear (ITS) and plastid (rps16 intron) DNA se- 
quences. 5S. African J. Bot. 70:407-416. 

Sun F.-J., S. R. Downie, and R.L. Hartman. 2004. An ITS-based phylogenetic analysis of the perennial, 
endemic Apiaceae subfamily Apioideae of western North America. Syst. Bot. 29:419-431. 

Sun F.-J.,G.A. Levin, and S.R. Downie. 2005. A multivariate analysis of Cymopterus glomeratus, formerly 
known as C. acaulis (Apiaceae). Rnodora 107:359-385. 

THEOBALD, W.L., C.C. Tsenc, and M.E. Matias. 1964. A revision of Aletes and Neoparrya (Umbelliferae). 
Brittonia 16:296-315. 


958 BRIT.ORG/SIDA 22(2) 


Book REVIEW 


HAKAN Rypinand Joun K. JecLum. 2006. The Biology of Peatlands. Oxford University Press 
(ISBN 0-19-852872-8, pbk.). Oxford University Press, 2001 Evans Road Cary, NC 27513, 
U.S.A. (Orders: 800-451-7556; http://www.oup.com/us/). $49.50, b/w figures and 
photographs, tables, graphs, and glossary, 343 pp., 61/4" x 91/4" 


The Biology ae tua nds by ae Rydin and oe is part of Oxford's Biology of Habitats series and offers 


readers a Peatlands. This is a well-illustrated book with many text- 
summarizing and research- eed illustrations. This book would be very suitable as a textbook for classes on 
peatlands or as a supplemental reading for exploring wetland habitats. Additionally, this title could also be a 


beneficial reference text a classes such as agronomy, ee or forestry. 
a 1) ¢. } 


The book < well alo oi ae on previous cha cere intorma- 
= 
tion. Chapter topics include (1) Peatl 


I | habitats, (2) at eee peatland 


habitat, (4) Sphagnum - the builder of boreal Jaen (5) Peat and organics soil, on re peat archives, (7) Peatland 
succession and development, (8) Hydrol peatlands, (9) Nutrients, lig )) Peatland pat- 
terns and landforms, (11) Peatlands Sau the world, (12) ae oe carbon bs nee ie oS) Uses, func- 


tions, and cue oncas of peatlands CI 


he text is very readable and is accompanied | by many supportive illustrations, graphs and charts. The 
authors’ eae chapters provide an introduction to terminology associated with peatlands which will help 
the reader both tear a are — ter Tminology and determine the relationship of peatland to 
} t 


other wetlands types. These openin 


the reader an overview of common plants and animals 
associated in peatlands as well as special sdepenions of these organisms to living in various peatland types. 


Chapter four focuses specifically on the life cycle, morp hology, physical attributes, chemical attributes and the 


colonization ability of Sphagnum mosses. 


All chapters are steeped with references to pertinent scientific studies. These references allow readers to 
learn more about methodology, data and results from such studies. The middle chapters of the book focus on 
peatland organic soils, hydrology including peat physical and chemical properties, peat as historical, fossil “ar- 


chives,” detailed explanations of the processes involved in peatland succession and devel t, descriptions of 


water flows, balances in peatland types and the roles of nutrients, light and appears in peatlands. The hy- 


drology coe includes a biguoats discussion of the variations in water chemistry along bog-rich fen gradi- 
Thi 


ents. from recent research studies on the subject and includes a summary 


chart of the surface water chemistry of various peatland sites. Within chapter nine is a wonderful example of 
research regarding forestry uses of peatlands in a discussion of “nutrients after drainage for forestry”; again the 
authors have presented many supportive research references in the discuss 

e final chapters discuss peatland patterns and formations “peatlands aed the world, peatland produc- 


tion n and catho balance, pec a and uses, peatland function and peatland management. These final chapters use the 
eatland ecology and ee The peatland 
landforms secon has a help ful inn on mire ste oue and c serene at different scales including 


eee ta able and detailed description of mire features, sites and types. Many aoe ert accom- 


pany this of mire types. The authors have focused this text on eas peatlands, but they do include 
sections contributed from any auLners on ana in Argentina, New Zealand, and southeast Asia. Peatland 


Ss acuen and a tie together much of what has been 
| | W icmp) i oli: : 1 fields. The pena: pues is loaded 
++ 1 a | } ) ] a ae ] 


W and various parts of the carbon cycle within peatlands, 


] 
biomass and rn Pp 
| 


as wellasa presentation of information on neat accum ae os limitations in accumulation “The nee chapter 
has many applications of lating susta lity and uses 


c 


in fields such as forestry, fuel use, horticulture a asa de- ies substrate. 
The Biology of Peatlands by authors Rydin and Jeglum provides a comprehensive overview to peatlands 
biology, ecology and peatland types. This mo) is Srull of supportive black and white illustrations, charts, tables 


an ndr ele erences to research studies. T Bi ls would set l textbook for classes focus- 


ing on peatlands, wetland habitat types, ind elated classes where peatland products may be of interest.—Lee 
Luckeydoo, Herbarium, Botanical Research Institute of lexas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 


SIDA 22(2): 958. 2006 


SIX NEW SPECIES OF SPHAGNUM (BRYOPHYTA: SPHAGNACEAE) 
FROM NORTH AMERICA 


Richard E. Andrus 
Environmental Studies sh 
Binghamton Univ 
Binghamton, New York 1 ne Ln U.S.A. 
randrus@binghamton.edu 
ABSTRACT 
Sphagnum mcqueenii, S. kenaiense, 5. bergianum, S. beothuk, S. sitchense, and S. talbotianum, are described 
and illustrated as new species for North America. 
RESUMEN 


raat | a ] N América Sp } S. kenaiense. S bergianum, 
1 


S. beothuk, S. sitchense, y S. talbotianum. 
INTRODUCTION 


Extensive field work in North America in recent years has revealed a number of new 
species of Sphagnum, mostly in more remote and poorly collected areas. Six of those spe- 
cies are described in this paper. Four are from Alaska, one from Newfoundland, one from 
both Newfoundland and Alaska and a sixth species from several states and Canadian 
provinces in northeastern North America. 


Sphagnum mcqueenii Andrus, sp. nov. (Figs. 1-6). Tyr: U.S.A. NEW HAMPSHIRE: Monroe Co. Town of 
Nottingham, Pawtuckaway State Park, Dead Pond, extensive poor fen, 18 Sep 1994, Andrus 9070 (HOLC 
TYPE: DUKE, ISOTYPES: BING, NY). Sphagnum sect. Cuspidata. 

Planta staturam mediam attingens, ate vel Devens escens, capitul pi pl ] i,g inali 

praedito. Cortex caulinuse i ietil litari it 

Folia caulina aequilaterali- triangularia, 0.75- te mm lata, 08> Ll mm Longa, late marginata, in Posen apical 

efibrillosa ie aballe osa pl | I I I iB 

] [Rey emecreed cee : | Tels li 


5 a | 7] 


praedita 
lanceola, 0.5- coe mm lata, ee 21 mm longa, in superficie convexa a cellulis hyalinus poros apicales O- i et 


L 


Ree i 12. 4-8 i diametro praedita. 
Plants robust and weak-stemmed,; yellow to light brown; capitulum typically flat- topped 
and with a + conspicuous terminal bud. Stems light green; cortex in 1-2 layers of moder- 
ately differentiated thin-walled cells. Stem leaves equilateral- triangular, 0.75-1.0 mm 
wide by 0.8-11mm long, usually spreading, apex obtuse, hyaline cells usually septate in 
lower half of leaf and efibrillose (or fibrillose near apex). Branches unranked, + straight, 
branch leaves moderately elongated at distal end. Branch leaves < 2.1 mm long, ovate- 
lanceolate, undulate and sharply recurved when dry; hyaline cells on convex surface with 
0-1 apical pores and often with pseudopores, concave surface with up to 12 round wall- 
thinnings in cell angles and sometimes along commissures; chlorophyllose cells trian- 
gular in transverse section and just enclosed on the concave surface. Sexuality unknown. 
Distribution.—Western Hemisphere Atlantic; known to date from Maine, New Hamp- 


SIDA 22(2): 959- 972. 2006 


BRIT.ORG/SIDA 22(2) 


Fis. 1. Sphagnum mcqueenii. Habit. 


ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 961 


Fic. 2. Sphagnum mcqueenii. Spreading branch. 


shire, Pennsylvania, and Vermont from the U.S.A. and Nova Scotia and Newfoundland in 
Canada. 

Identification—Sphagnum mcqueenii can be separated from S. cuspidatum Hoffm. 
by its enclosed chlorophyll cells, obtuse stem leaves and the relatively short, recurved 
and broad ( width: length ratio > 0.3) branch leaves. Sphagnum viride Flatberg. will dif- 
fer in having branch leaf chlorophyll cells not reaching the concave surface and pointed 
stem leaves. Both S. cuspidatum and S. viride are also smaller and slenderer plants. S. 
pulchrum (Braith.) Warnst. has strongly 5-ranked branch leaves, more pointed stem leaves 
and more well-enclosed branch leaf chlorophyll cells. Wet-growing forms of S. pulchrum 
will also not have branch leaves elongated at the distal end of the branches, a character- 
istic of the more “aquatic” Cuspidata species. The other “aquatic” Cuspidata with which 
S. mcqueenii can occur are S. torreyanum Sulland S.atlanticum Andrus. Both of these are 
larger plants and have longer branch leaves (> 2.1mm). They also have stem leaves more 
appressed to the stem than S. mcqueenii, in which the stem leaves are often spreading. 

Ecology.—Habitat preferences not well known but it has so far been collected in poor 
fen habitats with floating mats, wet carpets and wet peaty depressions. It was quite abun- 
dant in the New Hampshire type locality, where it occurred in the moat at the bog mar- 
gin associated with S. torreyanum, S. atlanticum, S. cuspidatum and S. flexuosum Dozy.& 
Molk 

Etymology.—The namesake of this species is the late Cyrus McQueen, a good friend 
and fellow sphagnophile, who led me to the type locality. 


Additional collections examined: CANADA. Newfoundland: N of Pouch Cove, Andrus 10309 (BING): Winterland, 
Andrus 10422, 10423, 10439 (BING, DUKE): N of Harbour Breton, Andrus 10632 (BING): Lark Harbour, Andrus 
10683 (BING, DUKE). NOVA SCOTIA. Guysborough Co.: Sc is & Maass 3315 (DUKE). U.S.A. MAINE: Acadia 
National Park, Little 16 (BING). Sagahadoc Co.: Allen 27060 (DUKE). PENNSYLVANIA. Monroe Co.: Andrus & 
Damman 6641 (BING). VERMONT. Windham Co.: McQueen s.n., i Nov 1998 (BING, DUKE). 


Sphagnum kenaiense Andrus, sp. NOV. (Figs. 7-12). Type: U.S.A. ALASKA. Kenai Peninsula Co. Kenai 
Peninsula, Soldotna, Headquarters Lake large poor fen complex surrounding lake, 17 Aug 2004 (HOLO- 
TYPE: DUKE; IsoTyPEs: BING, NY). Sphagnum sect. Cuspidata. 

Planta Denv caule debili, pallide brunnea vel aetate aurea, capitulo plano; cortex caulinus parietibus tenuibus, 

tantum leniter dilatatus. Folia caulina ad caulem appressa, lingulata, ovato-lingulata vel triangulata, aequa vel 

minora quam 0.9 mm longa, apice obtuso aus sata eroso ve ius see tantum leniter 5-ordinati vel 5- 


seriata, in apice distali rameali vix elongati. F 2-3 pendentibus 
compositus. Folia ramulina 1.1-1.3 mm longa, ovata, rigentia, in statu sicco ae antele atque subrecurva, 


962 BRIT.ORG/SIDA 22(2) 


Fics. 3-6. Sphagnum mcqueenii. 3. Branch leaves. 4. Stem leaves. 5. St 


} +] Veesenten “| a4. t lat ] 


cellulis hyalinis in parte mediana } s usque longitudinis 0.3- 
0.4) et in superficie convexa in tas folii intone saepe jaa oro magno atque/usque 6 pone ae et 
{ - ~ ¢ 


parte apical isae epe g i) 5 pe udoporis instructa, uperticie concava ad t 


E 
atque angulos areis parietum tenuium magnis rotundis Hee ee inconspicuuis vel nullis praedita; cellulae 
chlorophylligerae in section transversali triangulares, typice in superficie bene inclusae. 


Plants small.and weak-stemmed; pale brown to golden brown; capitulum flat-topped and 


ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 963 


12. Branch leaf 


Fig. 7-13 5} f 9 k j 7. Habit. 8. Branch. 9. Stem leaves. 10. Branch leaves. 11.B h leaf 
concave surface. 13. Stem section. Scale bars: A= 2 mm, 7; B = 2 mm, 8; C= 0.5 mm, 9-10; D = 50 pm, 11-13. 


only weakly 5-radiate. Stems pale yellow; stem cortex moderately well-differentiated but 
not much enlarged. Stem leaves appressed to stem; lingulate, ovate,to triangular; equal to 
or less than 0.9 mm long; apex obtuse and often erose to lacerate. Branches with leaves 
unranked to 5-ranked, leaves not much elongated at distal branch tip. Branch fascicles 
with 2 spreading and 2-3 pendent branches. Branch leaves ovate, 1.1-1.3 mm long, stiff, 


964 BRIT.ORG/SIDA 22(2) 


weakly undulate and slightly recurved when dry; hyaline cells in mid-region quite short 
and broad (width:length = 0.3-0.4), in lower 1/2 of leaf on convex surface often with 1 
large pore apically and/or up to 6 free pores, in apical region often with pseudopores 
along the cell margins; on concave surface with large round wall- Tenens in the cell 
ends and angles (these sometimes faint or absent); chlorophyllose cells triangular in trans- 
verse section and typically well-enclosed on concave surface. Sexuality unknown. 

Distribution.—South-Central and Western Alaska. 

Identification —Within its range, S. kenaiense may be confused with S. angustifolium 
(Russow) C. Jens., S. balticum (Russow) C. Jens.) and S. brevifolium (Braithw.) Roell. Sph- 
agnum angustifolium in the field has narrower ovate-lanceolate branch leaves. Micro- 
scopically S. angustifolium has a poorly differentiated stem cortex, chlorophyll cells not 
well-enclosed and narrower branch leaf hyaline cells. Sphagnum balticum has narrower 
branch leaves and typically only one hanging branch, with stem leaves markedly spread- 
ing. Microscopically the branch leaf hyaline cells are also narrower. Sphagnum brevifolium 
has stem leaves that are apiculate to acute and ovate-lanceolate branch leaves. It also has 
narrower hyaline cells in its branch leaves than S. kenaiense. 

Ecology.—Sphagnum kendiense occurs in poor to medium fens, often of a sedge domi- 
nated nature. It typically forms sprawling patches in hollows, often over bare peat. 

Etymology.—The species epithet kendiense is after the Kenai Peninsula where the 
species was found. 


a. 


Additional coll ined: U.S.A. ALASKA: King Salmon, Andrus 9306 (BING, DUKE); Selawik, Schofield 
121,733 (BING): Anc horage C ampbell Airstrip fen Andrus 8448 (BING, DUKE); Kenai Peninsula, Mil epost 100 on 
Seward Hwy., Andrus 8752 (BING, DUKE, NY). 


Sphagnum bergianum Andrus, Sp. NOV. (Figs. 13-19). Type: US.A. ALASKA. Kenai Peninsula Co: Kenai 
Peninsula, Soldotna, H leadquarters | ake, extensive poor fen con nplex at lake en 17 Aug 2004, Andrus 
10020 (HOLOTYPE: DUKE; ISOTYPEs: BING, NY). apa section Acutifolic 


pl] | a ] yb ] ] ] 
Ne 


omnino atro-fusco, 


St | 
centro Cz cenit aurato, in statu vivo saturate € rubritinge nti, in statu sicco nitore roseolo- pur Durascenti 
1: 


praedito.C aulis atro-fuscus Folia caulina late triangulari- lingul ata, 


1.15-1.25 x 0.8, in ae subobrusa vel late aiaicn limbo ad hasem ates oe cellulis hyalinis pro parte 


maxima |-septatis, [ vel isin parte mediana [ ami plus minusve 5-ordinati. 
Fasciculus see e cael: 2 patentibus atque | aie Folia ramulina 1.2-L3 x 0. 65-0.75 mm, late ovata, 
apice involuta, cellulis h yalinis in merosis rotundatis i ellipticis poris commissuras se- 
cus instructis, in apic idatim usque ab yasem maioribus sa, POTIS paucis 


magnis in epost dicate ibe paivasiint exceptis. Status sexualis ce Sporae 26-30 —- 


Plants moderate-sized to robust, capitulum more or less flat-topped and large; dark brown 
overall with a golden center to the capitulum and a distinctive deep red tinge in the field 
which becomes a pinkish-purple sheen upon drying. Stems dark brown; superficial cor- 
tical cells aporose. Stem leaves broadly triangular-lingulate, 1.15-1.25 = 0.8, apex slightly 
to broadly obtuse, border moderately broadened at base; hyaline cells mostly l-septate 
with afew to many 2 septate in the mid-region, shape rhomboidal. Branches more or less 
5 ranked. Branch fascicles with 2 spreading and | hanging branch. Branch leaves |.2-1.3 
mm x 0.65-0.75, broadly ovate, apex involute; hyaline cells on convex surface with nu- 
merous round to elliptic pores along the commissures, these grading from small pores at 
the apex to large pores at the base; concave surlace apor xcept for a few large pores in 
the lower side regions. Sexual condition dioicous. Sas 26-30 um. 
Distribution.—Newfoundland and South-central Alaska. 


ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 965 


Fics. 14-20. Sphagnum bergianum. 14. Habit. 15. Branch fascicle. 16. Stem leaves. 17. Stem leaf hyaline cells. 18. Branch leaves. 19. 


Branch leaf, concave surface. 20. Branch | face.A = 2 mm, 14; B = 2 mm, 15;C=0.5 mm, 16, 18; D = 50 pm, 17, 19-20. 


Identification. Sphagnum bergianum seems most clearly related to S. subfulvum 
Sjoers sensu lato, as described by Flatberg (1985). Both S. subfulvum and S. subfulvum ssp 
purpureum Flatb. are a lighter golden-brown than the quite dark brown of S. bergianum. 
Furthermore, both of these taxa have narrower stem leaves that are more acute at their 
apex. And neither of these taxa have distinctly 5-ranked branch leaves like S. bergianum. 
Sphagnum bergianum is a quite unusual looking species in the field, with its very dark 
brown color tinged with a deep red color. When dry this deep red becomes more purplish 


966 BRIT.ORG/SIDA 22(2) 


but it still shows some red. This distinctive field appearance made it quite recognizable 
when collected in Newfoundland in 2005 

Table 1 highlights the differences between S. bergianum and other larger brown 
Acutifolia. Because of its similarity, S. subfulvum ssp purpureum is included for complete- 
ness even though the author has not seen any North American specimens of that taxon 
and believes it to be of doubtful occurrence there. 

Ecology.—In the few sites where it has so far been collected, S. bergianum has been a 
hummock forming species of medium fen habitats, associated with such species as S. 
warnstorfii Warnst., S. subfulvum, S. platyphyllum (Braithw.) Warnst. and S. fuscum 
(Schimp.) H. Klinggr. 

Etymology.—This species named after Ed Berg, ecologist with the US Fish and Wild- 
lite Service, who first found this species in Headquarters Lake next to his office. 


Additional collections examined: CANADA. Newfoundland: 9 km N of Badger, Andrus 10646 (BING, DUKE, NY); 
15 km SW of Corner Brook, Andrus laa DUKE, NY). U.S.A. Alaska: Kenai Peninsula, Milepost 40 on 
Seward Hwy., Andrus 10036 (BING, DUKE 


a 


Sphagnum beothuk Andrus, sp. nov. (Figs. 20-25). Type: CANADA. NEWFOUNDLAND: Lark Harbour, 
ev, Andrus 10687 (HOLOTYPE: DUKE; ISOTYPE: 


Naas} 


ca. 50 km NW of Corner Brook, medium rich fen, 20 m e 
BING, NY) eee sect. Ac a id. 


Pl. anta parva | 1 | | fi usco nitore 1 li 


Caulis brunneus, eda corticalis supericlie eporosis. Folia oan ieee ll- 124 mm oe in apice 
subapiculata vel maximam partem lata, erosaque vel lacerata, me Ss ad co tantem parce dilarato, cellulis 


hyalinis rhombiformibus, 0-1-septatis. Rami plus minusve 5-ordin 2 patentibus 
atque 1 pendenti compositus. Folia ramulina 0.95-1.3 mm nae ovata vel ovato-lanceolata, concava, stricta vel 


parce en cecunee in pie volta, cellulis hyalinis in ie Hole convexa ee commissuras secus numerosis 


4 lell; | 
rot undatls ad 


5 \ L 


ie} 
s | titudinic Thal 


vel perpusillis on m, minoribus quam 025 par form ane superticie 


concava in partibus proximalibus ne poris parcis, rotundatis per sate instructa. Status sexualis 
ignotu. 

Plants small to moderate-sized; capitulum rounded and dense; dark brown with a pur- 
plish sheen. Stem brown, superficial cortical cells aporose. Stem leaves lingulate, 1.1-1.2 
mm long, apex slightly apiculate to mostly broad and erose to lacerate, border only 
slightly broadened at base; hyaline cells rhomboidal and 0-1 septate. Branches more or 
less 5-ranked. Branch fascicles with 2 spreading and 1 pendent branch. Branch leaves 
0.95-1.3 mm long, ovate to ovate-lanceolate, concave, straight to slightly subsecund, apex 
involute; hyaline cells on convex surface with numerous round to elliptic pores along the 
commissures, grading from large pores at the base to a mixture of small and tiny (2 pm, 
less than 0.25 cell width) at the apex, concave surfacewith a few large, round pores/cell 
in lower side regions. Sexual condition unknown. 

Distribution —Known only from Newfoundland, Canada. 

Identification.—In the field S. beothuk looks like a very dark form of S. fuscumexcept 
for its distinct purplish sheen. It’s rather more robust than Sfu scumand also has 5- Sans 
branch leaves, which S.fuscum usually lacks. Microscopically, however, S. beothuk differ 
clearly in the tiny pores seen mixed in with more typical pores on the convex euiiace of 
the branch leaf at the apex. Among the known Acutifolia species, only the red pigmented 
S. warnstorfii has been noted with this character before. 

Ecology.—Sphagnum beothuk forms dense hummocks similar to S. fuscum. It appears 
to be a minerotrophic species and has been collected in the same mire as Ssubfulvum, S. 
inundatum Russow, and S. warnstorfii. 


Taait 1. Comparison of Sphagnum bergianum and related species. 


Characteristic 


5. bergianum 


S$. subfulvum 


$. subfulvum ssp. 
purpureum 


5. subnitens 


5. flavicomans 


Color 


Branch fascicles, 

spreading + hanging 
branches 

Branch leaf ranking 

Branch leaf shape 

Stem leaf shape 


Stem leaf apex 


Stem leaf length in mm 


Stem leaf hyaline cells 
Distribution 


Dark brown with 
a deep red tinge 


Often 5 -ranked 
Broadly ovate 
Triangular-lingulate 


More or less obtuse 
.15-1.25 

|—2 septate 
Newfoundland & 
SC Alaska 


Golden-brown with a 
purplish gloss sometimes 
2+ 


Unranked 

Ovate 
Triangular-lingulate to 
broadly lingulate 
More or less acute 
ee 

0-1 septate 

NE and NW North 
America 


Golden-brown with 
red tinge 

+ 1-2 
Unranked 

vate 
Triangular-lingulate 
More or less acute 
1.3-1.4 


O-1 septate 
NW Europe 


Reddish-purple mixed with 


golden brown 
2+ 1-2 


Unranked 


Ovate to ovate-lanceolate 


Triangular to triangular- 
lingulate 

Acute to apiculate 
1.3-2.7 

0-1 septate 

Eurasia, W North America 
& New Zealand 


Brown 


2+ 1-2 


Often 5-ranked 
Ovate-lanceolate 
fail triangular- 
lingulat 

Acute to ice 


0-1 septate 
NE North America 


VOIYIWY HLYON WOUd WANOWHdS 40 S31D3dS MIN ‘SNYGNY 


196 


968 BRIT.ORG/SIDA 22(2) 


ABCD 


1D 


| AY 


Op 
Oe 
C) 

EO 


o 


/ 


ae 


gua7cee. 
i isc qs 
WANS 

exe 7 


rO) 


i 


| 


Fics. 21-26. 5; k. 21. Habit. 22. Branch. 23. Stem leaves. 24. St | Ils. 25. Branch leaves. 26. Branch leaf 


hyaline cells, convex surface. A = 2 mm, 21; B = 1 mm, 22; C= 0.5 mm, 23, 25; D = 30 pm, 24, 26. 


Etymology.—The Beothuk are the aboriginal people of Newfoundland. 

Comments.—Several things suggest a possible hybrid origin between S.fuscum and S. 
warnstorfii. The tiny pores at the branch leaf tips, the purplish sheen and the 5-ranked 
branch leaves suggest S. warnstorfii and the lingulate stem leaves are a character of S. 
fuscum. Furthermore the type locality is a minerotrophic site in which both S. fuscum 
and S. warnstorfi also occur. 


ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 969 


YG; my it pia 
(i dart LE pe 
A KA 
SSE 
" Vy we 


Fics. 27-34. Sph itch 27. Habit. 28. Branch. 29. Branch leaves (“b”) 30. Stem with leaves. 31. St leaf hvall IIs. 32 
£ 24 RQ hl £ 


A=2mm,27;B=1mm,28, 30;C=0.5 mm, 29, 32; 


hl 


Stem leaves (“s”).33.B 
D=50 pm, 31, 33, 34. 


Additional collections examined: CANADA. Newfoundland: Lark Harbour, Andrus 10685, 10686, 10687 (1orotyPEs: 
BING, DUKE); ca. 16 km S$ of jet. of Hwy 210 and Hwy 214, Andrus 10408 (BING, DUKE). 


Sphagnum sitchense Andrus, sp. nov. (Figs. 26-33). Type: US.A. ALASKA: Sitka Co. Sitka, Harbor Mt, 
alpine area at end of road, 700 m elev, dense patch in tundra, 19 Aug 2004, Andrus 10097 (HOLOTYPE: 
DUKE, IsoTyPEs: BING, NY). Sphagnum sect. Acutifolia. 


970 BRIT.ORG/SIDA 22(2) 


nee F eee F ] dence. loat lo ni lesti Caulis roseolus 


. | c Ct 
ediis coneaioue fF aialibie p — Folia caulina anguste vane aria vel lingulato-triangularis, 1.2- 


ven mm longs x 2 6- 2 8) mm oe ratione enihtieticg ongicncinls usque SAUCES ca. 2:1, apice apiculato, limbo 
ellulis lum 2-septatis, saepe in 


apice kenbillods Bata non 5-ordinati. Fasciculus ramulinus e scene? 2 patentibus atque | absaiae peepee 
aes ramulina 1 = Wo mm longa, 0.5- ve 55 mm lata, ovato-lanceolata, concava, stricta, in 


l 
vel ellipticis ad basem poris magnis 


jen 


Bree a vege ad ee rotunealls quoae staturam mediocribus, in superficie concava poris magnis 


rotundatis g 

Plants cael saints flat-topped and dense; pale yellow and pink, lacking 
sheen. Stem pink, superficial cortical cells aporose. Stem leaves narrowly triangular to 
lingulate- triangular, 1.2-1.6 mm long x 0.6-0.8 mm wide, length:width ratio ca 2:1 apex 
apiculate, border strong and moderately broadened basally; hyaline cell rowly rhom- 
boidal and | to occasionally 2 septate, often fibrillose apically. Branches not 5-ranked. 
Branch fascicles with 2 spreading and | pendent branch. Branch leaves 1.3-1.5 mm long 
and 0.5-0.55mm wide, ovate-lanceolate, concave, straight, apex involute; hyaline cells on 
convex surface with numerous round to elliptic pores along the commissures, grading 
from large elliptical pores at the base to moderate-sized round pores at the apex, concave 
surface with a few large round pores scattered throughout. Sexual condition unknown. 

Distribution—Known only from the type locality in pe neeee Alaska but since al- 
pine areas in this area are poorly collected it is likely more widespr 

Identification Sphagnum sitchense is somewhat similar to S. cari ilyeinen (Ehrh.) 
Hedw. But differs ina number of important ways. Its stem leaves are often narrowly tri- 
angular whereas those of S. capillifolium are lingulate- triangular. Branch fascicles on S. 
sitchense have 2 spreading and one hanging branch whereas those of S. capillifolium have 
2 spreading and mostly 2 hanging branches. The stem leaf hyaline cells in S. sitchense 
are 1-2 septate while those on S. capillifolium are 0-1 septate. In the field S. sitchense has 
a pale red tinge and quite flat topped capitulum which contrast with the deeper red tinge 
and rounded capitulum of S. capillifoilum. 

Sphagnum sitchense also bears some resemblance to Sphagnum subnitens but differs 
in the following ways. Sphagnum sitchense hasa strongly bordered stem leaf with afew 2 
septate hyaline cells while S. subnitens has a narrow border and hyaline cells 0-1 septate 
only. Sphagnum sitchense also lacks the metallic sheen and brown coloration that are 
typical of S. subnitens. And lastly, the branch fascicles in S. sitchense have only one hang- 
ing branch whereas S. subnitens has mostly two. 

The stem leaves of S. sitchense are quite similar to those of S. quinquefarium, with 
which it occurred in the type locality, but the latter species has 5-ranked branch leaves 
and typically 3 spreading branches per fascicle. 

Ecology.—In the type locality, S. sitchense was collected several times in dense patches 
in the tundra. Growing similarly in the same area were S. girgensohnii Russow, S. 
quinquefarium (Braithw.) Warnst., and S. compactum D.C. 

Etymology.—Sphagnum sitchense is after the type locality. 


— 


Additional collections examined: U.S.A. Alaska: Sitka, Harbor Mt,. Andrus 10095, 10098 (BING). 

Sphagnum talbotianum Andrus, sp. nov. (Figs. 34-39). Type: U.S.A. ALASKA: Bethel Co: Bethel, 0.9 
mi — es Rd. ie turnoff for Chief Eddie Hoffman Hwy, just W of Larson Subdivision, medium fen, 

=; DUKE; ISOTYPES: BING, NY, TRH). Sphagnum sect. ea 


Planta rubri-pigmentifera, capitulo in apice complanato, subdenso. Cortex caulinus eporosus. Folia caulina 


ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 971 


ABCD 


Fics. 35-40. Sphag lboti 35. Habit. 36. Two branches. 37. Stem leaves. 38. Stem | 
Branch leaf, convex surface. A = 2 mm, 35; B = 2 mm, 36;C = 0.5 mm, 37, 39; D = 30 pm, 38, 40. 


ve obtusa, 0.8-1.05 mm longa, 0-0.55 mm lata, limbo ad 
0-1 septatis,et fibrillis et poris carentibus. Folia ramulina 
= j; ] ; lselasss falas 


lingulato-triangularia vel lingulata, in apice plusminus 
basem valde distincto, cellulis hyalinis rl bif il 


stricta et ovato lanceolata, 1.0-1.2 mm longa, 0.3-0.35 mm lata 


t k = a, 
convexa poris quoad amplitudinem mediocribus, in apice 4-6 um usque in base minoribus quam 20 pm, in 
ee ; eee 7 


] 1st 


superficie concava ef ae praeter interdum in partibus I g I 
Status sexualis ignotis. 

Plants pink-colored; capitulum flat-topped and moderately dense. Stem red-tinged, cor- 
tex eporose. Stem leaves triangular-lingulate to lingulate with a more or less obtuse apex, 


972 BRIT.ORG/SIDA 22(2) 


0.8-1.05 mm. long = 0.5-0.55 mm. wide; border strongly developed at the base; hyaline 
cells rhomboidal and 0-1 septate, efibrillose and eporose. Branches with leaves 5-ranked 
and loosely spreading. Branch leaves small, 1,0-1.2 mm. long « 0.3-0.35 mm. wide; con- 
vex surface with moderate- sized pores, grading from 4-6 jim near apex to more than 20 
jum at base, concave surface eporose except for occasional large round pores in proximal 
side regions. Sexual condition unknown. 

Distribution.—Western Alaska. Currently known from Bethel, King Salmon, and 
Koyukuk National Wildlife Refuge. 

Identification.—As far as other red Acutifolia within its general range, S. talbotianum 
is most similar to S. rubellum Wils. and S. warnstorfii. Sphagnum rubellum has a lingulate 
stem leaf with at least some hyaline cells 2-septate whereas S. talbotianum has stem leaves 
that are often triangular-lingulate and with hyaline cells that are 0-1 septate. The branch 
leaves on S. rubellum are also often subsecund while those of S. talbotianum are straight. 
Sphagnum warnstorfii looks quite similar macroscopically but has much smaller ringed 
pores on the convex surface of the branch leaf tips. 

Ecology.—Collections to date indicate this to be a quite widespread tundra species of 
minerotrophic peatlands. It typically forms low dense hummocks. Compared to 
S.rubellum Wils. and S. warnstorfii Russow, the other abundant red quinquefarious 
Acutifolia with which its range overlaps, S. talbotianum occurs in richer sites than S. 
rubellum and poorer sites than S. warnstorfii.. Common associated Sphagna include S. 
lenense Pohle. S. squarrosum Crome, S. teres (Schimp.) Aongs., S. brevifolium (Braithw.) 
Warnst., S. arcticum Flatberg & Frisvoll, S. subsecundum Nees, S. perfoliatum Savicz- 
Lubitskaya, S. obtusum Warnst., S.alaskense Andrus & Janssens, and S. steerei Andrus. 

Etymology.—Sphagnum talbotianum honors Stephen Talbot, who has collected more 
Sphagnum in more remote areas of Alaska than any other botanist. He deserves more 
credit than anyone else for our current knowledge of the distribution of Sphagnum in 
that state. 


Additional coll ined: ALASKA: Koyukuk National Wildlife Refuge, 21 Aug 1989, Tal bot Talbot 8-8- 
16 (BING); Yukon Delta National Wildlife Refuge, Kuskokwim River Delta, (mixed with S. lenense) 15 Aug 2001, 
Andrus 9465 (BING, DUKE, NY); King Salmon, near village, 5 Jul 2001, Andrus 9522 (BING, DU “ NY); King 
Salmon area, ca. 39 km SW on maritime tundra, 6 Aug 2001, Andrus 9354 (BING, DUKE, NY). 

REPERENCE 


FLATBERG, KI. Studies in Sphagnum subfulvum Sjoers, and related morphotypes. Lindbergia 11:38—-54. 


REVIEW OF CRATAEGUS SERIES PULCHERRIMAE (ROSACEAE) 


J.B. Phipps RJ.O’Kennon 
Department of Biolog Botanical Research Institute of Texas 
The University of Western Ontario 509 Pecan St. 
London, Ontario, N6A 5B7, CANADA Fort Worth, Texas, 76102-4060, U.S.A. 
jphipps@uwo.ca okennon@brit.org 
KA. Dvorsky 
Dept. of Biology 


The University of Western Ontario 
London, Ontario, N6A 5B7, CANADA 


ABSTRACT 


Crataegus series Pulcherrimae (Rosaceae) is reviewed, Beadle’s group Sargentianae being incorporated. Ten spe- 
cies, C. sargentii, C. gilva,C. mendosa, C. pulcherrima, C. pinetorum, C. venusta, C. opima, C. tecta, C. pallens, and C. 


eximid are recognized. In addition, a possible eleventh species, C. ?vobur, is suggested but the evidence is not yet 


final on this. All species are lectotypified here except C. pinetorum, which is neotypified. Epitypes are proposed 


for two species, C.opima and C. venusta. Synonyms are also typified. Each species is keyed out and is represented 
by the most detailed description to date. Detailed line illustrations and county level range maps, in both cases 
generally the first for each species, are provided. Repesentative specimens are cited. 


Key Worps: Crataegus, series Pulcherrimae, group Sargentianae, typification 
RESUMEN 


Se revisa Crataegus serie Pulcherrimae (Rosaceae), se incorpora el grupo ma eke nae de Beadle. Se reconocen 


diez especies, C. sargentii, C. gilva,C. mendosa,C. pulcherrima,C. pinetorum,C. venusta C.opima, C. tect pallen 
y C eximia. demas se sugiere una posible undécima So oe cad ee no hay pruebas finale aun. Se 
lectotipifican todas las ee es C. pinetorum 


£ c c [ A ft Ms 


. ona venus iste los sinonimos. Se aport 

llada hasta la se S li il i detalladas y | 
L Lt > i 
] 


1d Aantiacad Lh ] 4 d ie S 
a) Lr I r r I 


if ] 
y of OLICCE la 
pera | ve 


The group Pulcherrimae, without formal rank or description, was first used by Beadle 
(1902) in the preparatory paper for his treatment in Small’s (1903) Flora of the southeast- 
ern United States, edition 2. In the two publications Beadle’s treatments are identical and 
itis notable that all but the Type species, C. pulcherrima, were described by Beadle. Beadle’s 
Pulcherrimae, which he treated as a group of nine species, was noticed by Sargent (1905) 
and was used in Palmer's (1925) nomenclator but it was not formally recognized until 
Robertson (1974) carefully validated the name. Beadle (1902) also recognized another 
group, Sargentianae, of 15 species, all described by him, that in my opinion cannot mean- 
ingfully be distinguished from Pulcherrimae and it is therefore here included with the 
latter group such that it now has 24 described species. 

Later, Tidestrom, in Small’s Manual of the Southeastern Flora (1933), dispensed with 
all the species of Pulcherrimae and Sargentianae, and henceforth floras of the region 
usually followed this example. A few, however, like Kurz and Godfrey (1962) did recog- 
nize the existence of the series and thus, a handful of its species continued to appear 
sporadically. 

This paper, based on the examination of 374 herbarium sheets from 25 herbaria and 
numerous field trips to the relevant region, reviews series Pulcherrimae as a whole for 


SIDA 22(2): 973-1007. 2006 


974 BRIT.ORG/SIDA 22(2) 


the first time since Beadle inaugurated the groups Pulcherrimae and Sargentianae in 
1902. Like other members of this series of papers, this one is a precursor to the author’s 
treatment in Flora of North America, vol. 9, due to be published next year. As such, it is 
not intended to make a deep analysis of serial boundaries or relationships but rather has 
the principal aim of assessing the individual species to be recognized. Nevertheless, a few 
obviously necessary adjustments to serial boundaries are provided in papers for this se- 
ries. Accordingly, lam characterizing series Pulcherrimae as being almost completely 
glabrous at maturity, having possession of 20 stamens, red or yellow fruit and a some- 
what elevated fruiting calyx. Though this diagnosis now includes Beadle’s group 
Sargentianae the expanded series does not appear to meaningfully widen the original 
diagnosis for ser. Pulcherrimae. The relationships of ser. Pulcherrimae are clearly to ser. 
Intricatae. Indeed, ser. Intricatae,a mainly 10-stamen group with a number of very hairy 
species, is not in other respects very different from ser. Pulcherrimae and the two may 
eventually be fused, a procedure which is, however, not appropriate to this paper. Series 
Pulcherrimae, whether considered in its narrow sense or in the extended sense (includ- 
ing Sargentianae), is restricted to the southeastern United States except for some over- 
flow into southeastern Texas. 

The approach used for this work, after the series boundaries had been settled on, was 
to study the maximum number of specimens possible over an approximate 15-year time 
frame. Accordingly, loans were requested from over 40 herbaria in the southeast plus 
several national herbaria with significant Crataegus collections. Only 24 institutions, 
however, were able to produce specimens of this series. Therefore the first author, assisted 
often by O’Kennon, made numerous field trips to the region to search for further speci- 
mens of ser. Pulcherrimae as well as other Crataegus species. The results of this field- 
work proved very helpful and review of the appendix of cited specimens shows that UWO 
now has more Pulcherrimae exemplars than the totality of specimens received on loan 
from all other herbaria. The specimens received on loan plus the UWO collections were 
then grouped into the more distinctive and repeatedly encountered morphotypes with 
few if any intermediates between them. These were then described in detail, where pos- 
sible, by matching with type material, which is well represented for this series. The names 
used were typified, a key to taxa recognized made and then line illustrations and distri- 
bution maps were produced, nearly always for the first time, for nearly all of the species 
recognized. The taxonomic conclusions are those of the first author while the county 
distribution maps were produced by K.A. Dvorsky, working from the UWO database of 
specimens. 

Difficulties working with series Pulcherrimae stem from certain problems in work- 
ing with Beadle’s text, particularly his imprecise description of leaf shapes as exempli- 
fied by his use of the phrase “leaf margins lobed and incised, leaves ovate” (or very simi- 
lar) for nearly all taxa, although inspection nevertheless demonstrates some clear and 
striking differences in leaf shape. Indeed, this shape characteristic proves to be the main 
taxonomic discriminator between species as recognized in this paper, rather than Beadle’s 
emphasis on fruit colour (red and yellow being usual in the series), fresh anther color, or 
flower size which do not seem to correlate sufficiently well with other characters to be 
generally useful. Furthermore, most collectors, including Beadle, have not recorded color 
characters with their label data, making them especially difficult to use with herbarium 
material. UWO collections are among the few that routinely record these characters. 
Another problem is that series Pulcherrimae does not seem to be particularly common 


— 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 975 


except in sporadic favored locations. It is difficult to tell how much more abundant mem- 
bers of the series were in Beadle’s time, but, although a number of label annotations indi- 
cate very high abundances, ie., ‘thousands’ (very unusual today), the main collectors of 
this period, Beadle himself and Harbison, Biltmore’s collector, collected few specimens that 
survive other than the early collections associated with the protologues. Thus it is essen- 
tially impossible to infer the level of change in frequency over the last 100 years or so. 

Apomixis, polyploidy and perhaps hybridization (Phipps, 2005) contribute to taxo- 
nomic difficulty in Crataegus but so far no one has studied breeding systems in ser. 
Pulcherrimae, and only two chromosome counts or ploidy level estimates exist. Talent 
and Dickinson (2005) report a triploid ploidy level from an unidentified species and 
Longley (1924) a triploid count from C. pinetorum. One cannot generalize from so little 
information. Hybridization is first suggested by morphological intermediacy and as such 
there is no definite suggestion of it occurring in ser. Pulcherrimae. 

This is, therefore, a somewhat provisional treatment of ser, Pulcherrimae and be- 
cause of the problems noted it is necessarily grounded in the work of Beadle, the only 
previous worker who has paid serious attention to the group. In attempting to place all of 
Beadle’s 1902 and 1903 names in his groups Pulcherrimae and Sargentianae into the syn- 
onymy for this paper I have occasionally come up with only a possible, rather than a 
definite match. This itself speaks to difficulties of species level synonymy in series 
Pulcherrimae. Further, whereas most specimens can be more or less straightforwardly 
accomodated in the taxa ‘a that Irecognize here, there is a general residue of about 3% (see 
Appendix of Cited S ) of specimens of adequate quality where this is less straight- 
forward. Because of ‘this somewhat provisional state of the taxonomy, we also provide a 
map to the collective distribution of the series (Fig. 1). Thus, this will not be the last word 
on ser. Pulcherrimae and even more collecting as well as some combination of morpho- 
metric work, biosystematic investigation and molecular studies will be needed to stabi- 
lize the understanding of all its species. It is therefore to be hoped that a competent sys- 
tematist, preferably conveniently based in the southeast, will be encouraged by the 
platform presented here to carry out such work. 

This paper is a continuation of the first author’s studies of Crataegus in the south- 
eastern United States, examples of which are Phipps (1988) and Phipps and Dvorsky 
(2006). Because these papers were originally intended to provide the detailed backup for 
the first author's treatment of Crataegus in the southestern United States for Cronquist’s 
now defunct flora of that area, our meaning of “southeastern United States” here is iden- 
tical to Cronquist’s, ie. Louisiana and Arkansas are the western limits for that flora. Texan 
records are, however, added but only because a few species are of sporadic occurence in 
the extreme southeast of that state. 

The text continues with the formal taxonomic presentation. 


TAXONOMY 
Series Pulcherrimae Beadle ex K.R. Robertson, J. Arnold Arbor. 35: 628. ate TYPE spect 
oe 


sae oneal Ashe, Pulcherrimae Beadle, group without rank or des Bot 
1:66. 1 Pulcherrimae Beadle ex cnet in clavem, Man. Trees North Amer 1:400. 1905; group 
ens EJ. Palmer, in clavem, J. Arnold Arbor. 5:78. 1925. 

Note.—Robertson attributed sectional rank for this group by Palmer (1925), but I argue that Palmer was not 
formally using the rank section there (Phipps, Harvard Pap. Bot., submitted) 


Sargentianae Beadle, group without rank or description, Biltmore Bot. Stud. 1:59.1902. 


976 BRIT.ORG/SIDA 22(2) 


oN Sak. SE e 
ra are yA a oe 
yt x LY 

rar — 


as 
gs 


ese PT | eS asiar ay 
Tree RES 
ioasemmmen <8 


ry 
ny, 


im 


pe a. gel 
= ‘ie 
ie a 


a, 


Shrubs or small trees; l-year old twigs shiny golden-tan to shiny deep reddish-brown; 2- 
yr old twigs deep gray; thorns mostly 2-3 cm long, fine, very dark at 2 yr. Leaves decidu- 
ous, long-petiolate; petiole glabrous, bearing a few to several, someti stipitate, glands; 
blades 4-7 cm long, ovate to ovate-rhombic or narrow-ovate in general outline, unlobed 
or more frequently with 4-6 shallow lobes per side, sometimes all of the lobes or only the 
upper, obscure, lobe tips blunt to acute; venation craspedodromous, 4-1] veins per side; + 
glabrous. Inflorescences 5-10 flowered slightly domed panicles; branches glabrous, bear- 
ing often plentiful, caducous, membranous, linear, gland-margined bracteoles. Flowers 
14-18 mm diam.; hypanthium externally glabrous; calyx-lobes narrow-triangular, glan- 
dular-serrate to nearly entire; petals + circular, white; stamens 20, anthers usually pink 
to purple, sometimes cream (e.g. C. venusta); styles 2-5. Fruit 5(Beadle)-13 mm thick, py- 
riform to + orbicular, glabrous, yellow, orange, ruddy, red, red-purple or green, calyx lobes 
usually reflexed on a short collar; pyrenes 2-5. Mature bark deeply corrugated at least in 
some. 

Series Pulcherrimae occurs from the Gulf Coast of eastern Texas to northern Florida. 
Its species are collectively fairly common over much of Mississippi and Alabama as well 
as the western part of Georgia. There are also sporadic records east to South Carolina, one 
record in southern Tennessee and one locality for North Carolina in Buncombe County. 
Ten definite and one possible species are recognized here. 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 977 


For the whole series, flowering is in April through much of Mississippi, Alabama 
and Georgia but, in panhandle Florida, it may start late March and in North Carolina or 
Tennessee in May. According to the season, typical timing may be retarded or speeded up 
by cooler or warmer weather than normal. The south-north progression is typical for 
spring flowering plants in the region (e.g. Reader et al. 1974). Fruit is ripe anywhere from 
late August to early October and its retention past the early stages depends on the mesicity 
of the site and seasonal weather until full ripeness makes the fruit attractive to frugivores. 

Related species of Crataegus usually show little or no habitat variation in their physi- 
cal habitat such that niche differentiation, if it exists, must be based on other character- 
istics. Series Pulcherrimae is normative for the genus in this respect as far as present ob- 
servation indicates and the plants are woodland edge or woodland species of mesic 
localities, if the latter, usually under oak or oak-mixed canopy but not in the denser shade. 
Soil type preference for the mesomorphic members of the genus is also not normally pro- 
nounced with the exception of avoiding coarse-grained or more acid soils. Species of ser 
Pulcherrimae are mainly recorded on the finer-grained soils. A reviewer makes the sug- 
gestion that the distribution of C. venusta, however, may be restricted to blackland soils,a 
soil type that certainly harbors hawthorns, but this cannot be confirmed from data avail- 
able to me. Three of the mechanisms for niche differentiation in Crataegus are: differen- 
tial anthesis timing, different pollinator preference and fruit differentiation (color, time 
of ripening, size, perhaps taste) and several of these differences do occur with ser. 
Pulcherrimae but have not been investigated further. Differences in anthesis time in sym- 
patric Crataegus may be very large, up to six weeks, but usually correlate to taxononomic 
distance (for more on this see Phipps 1995, on hybridization), so they likely do not ini- 
tiate niche differentiation. In the case of ser. Pulcherrimae different taxa that occur at the 
same site do sometimes have slightly different, but overlapping, anthesis time but there 
is no consistent recording of this. There is also an untested suggestion that differences in 
fresh anther color (basically, anthocyanic or not) may affect pollinator faithfulness, thus 
beginning niche dif iation. In this respect it is interesting that such color differences 
quite often appear in very closely related taxa, though they are not particularly frequent 
within single species. There is a huge field of study here awaiting attention by an enter- 
prising biologist but it clearly needs preceeding by a plausible taxonomy against which 
it can be tested. 

The deeply corrugated bark of several species, found in few other series, glabrous or 
nearly glabrous plant parts (except in one aberrant specimen), about 20 stamens, rather 
glandular-bracteolate inflorescences, usually spreading, somewhat elevated calyx-lobes 
in fruit, and sometimes large numbers of lateral veins in the leaf make this series distinc- 
tive. It is thus a quite easily recognized series, except for its differentiation from ser. 
Intricatae, which latter normally possesses 10, not 20, stamens as in ser. Pulcherrimae 
and has some very hairy species. As in series Intricatae, yellowish-fruited forms are at 
least as numerous as red-fruited ones. 

Notes on terminology: 

1. This paper uses the term ‘leaf incision index, abbreviated to LII, for the relative 
depth of sinuses. This is measured by joining the lobe tips and projecting a line 
from there through the deepest part of the sinus, parallel to the adjacent veins, to 
the mid-vein. A one hundred percent value indicates cut through to the mid-vein 
50% cut half-way, cut 25% of the way to the mid-vein 

2. “Leaf? always refers to short-shoot leaves unless indicated otherwise. 


978 BRIT.ORG/SIDA 22(2) 


KEY TO SERIES PULCHERRIMAE 


A He tara branches glabrous [series Pulcherrimae, sens. str.]. 
. Well-developed short-shoot leaf blades fairly narrow, 1.5-2 * as long as broad. 
2. Well-developed short-shoot leaf-blades narrow-elliptic to narrow-ovate in general out- 
line;lobes absent or very shallow, perh ne notches with Ll 5% or even 10%, but if so 
| regularly occurring in all or most eves ee veins 6-10/side. 
. Leaf blades broad-elliptic to narrow-ovate in general shape; apices subacute to blunt; 


margins with crenate or crenate-serrate teeth; fruit red 3.C. mendosa 
3. Leaf blades narrow ovate to lanceolate in general shape; apices acute to acuminate; 
arginal teeth sharply acute; fruit yellowish 2.C.gilva 


2. Well-developed short-shoot leaf blades broad-elliptic or broad-oblong to narrow-ovate 
in general outline; regularly and distinctly lobed; max. LIl 10-40%; main lateral veins 4—6/ 


side. 
4. Lobes 2-3 per side, blunt, max. LI <15% 4.C. pulcherrima 
4. Lobes 2-5 per side, blunt or acute, max. LIl <15—25% or more. 

5. Leaf blades narrowly elliptic to very narrow-ovate in general outline; 1-2 (—3) shal- 


low, acute lobes per side, max. LI usually <15% 5.C. pinetorum 
5. Leaf blades narrowly ovate in general outline; 4-5 deep, acute to subacute, usually 
narrow-cuspidate lobes per side, max. LI] often 25% or more 9.C. incilis 


1. Well-developed short-shoot leaf blades broader in proportion, generally < 1.5% as long as 
wide 


6. Many leaves of a distinctive type, + trullate-ovate in general shape, the terminal half + 
straight-sided across lobe-tips; 4-6 cm long; lobes 1—3/side, sharp, but very small to small 
(max. LI] 2-10%(-15%)); 5-7 lateral veins/side; anthers pink; fruit yellowish 1.C. sargentii 

6. Leaves of quite different construction; if with few, short, very acute lobes then blades 
more ovate or oblong in form and the terminal half not + straight-sided across the lobe 
tips; length various; lobes nearly always much more pronounced, at least at ae ve- 
nation various; anthers ivory to cream or pink to purple; fruit yellowish or reddis 
7. nae with well-marked lobes, max. LI] usually 15-25%, ee asia more. 

. Blades usually smaller, 3—5(—8) cm long, ovate to broad g 
angular or not; anthers and fruit various colors. 
9. Majority of leaves broad-ovate in general on with 4—5 lateral veins per side 
and 3-4 lobes;lobes short or longer (max. LIl 10-40%). 


10. Lobes + blunt, generally fairly short (m3max. LI] 20-40%) 8.C. tecta 

9. Majority of leaves ovate in general shape with 5-8 lateral veins per side and 4—- 
6(—7) lobes; lobes fairly short, OOIUEP to acute (max. LII 10-40%) 7a.C. cf.opima 

8. Blades Be 5-8 cm long, rhombovate in general shape; lobes very angular;anthers 


10. C. eximia 


cream; fruit red 
7. Leaf pede: Hn alot lobes, max LI] 5— 10%. 
11. | quite shar 
eaf blades ee oblong to ei ovate; lobes and teeth acuminate iim? 
evelopment, sometimes reflex 6.C. venusta 
12. Leaf blades narrow ovate to ovate-oblong; tips of lobes and marginal teeth 
less pronounced. 
13. Leaf blades ovate-oblong in general shape; angle at leaf- — Aas ca. 90° 
shee — form 
13, ou blades narrow-ovate in general shape; angle at leaf- ‘ne ae 60° 


2.C.gilva 
11. Lobes cn to subacute. 
14. Leaf blades generally large at maturity, mostly 5-8 cm long; lateral veins 5-7 
per side; blades nearly unlobed to shallowly lobed; bases cuneate 3a.C.cf. 
C. mendosa 


14. Leaf blades small to medium-sized, rarely many > 5 cm long; lateral veins 4—7 
per side; blades regularly lobed, but sometimes shallowly; bases cuneate to 
rounded. 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 979 


15. Leaf blades cuneate at the base, most blades approximately 1.5:1 4.C. 

pulcherrima 

Leaf blades broad-cuneate to rounded at the base, blades <1.5:1 .C.opima 

B. Inflorescence branches pubescent C. craytonii ser. Intricatae] 


1. Crataegus sargentii Beadle, Bot. Gaz. 28:407. 1899. (Fig. 2.) Typr: U.S.A. ALABAMA. De Kalb Co: 
ial Head, 20 — 1899, C.D. Beadle 1289(LECTOTYPE selected here: US 969465; ISOLECTOTYPE: US 969466). 
2=C ili {le, Biltmore Bot. Stud., 1:68. 1902. Type: U.S.A. FLORIDA. Jackson Co:Chattahoochee, 

8 Apr 1901, TG. Harbison H4096 (LECTOTYPE selected here: US 969401). 


Shrubs or small trees 3-5 m tall; extending twigs glabrous, reddish; at 1 yr. shiny deep 
reddish-brown, 2 yrs grayish-red, older gray; thorns 2-5 cm long, few to numerous, slen- 
der, straight to slightly recurved, black at 2 yrs. Leaves: petioles 25-40% length of blade, 
slender, glabrous, with a few sessile glands; blades 4-6(-7) cm long, ovate-trullate in gen- 
eral outline; tip acute to acuminate, base rounded to broad cuneate; sides barely or shal- 
lowly 1-3 lobed; lobes acute, max. LII usually 0-10%; upper third of leaf often unlobed; 
margins finely and sharply serrate with numerous very small teeth usually <1 mm long: 
venation craspedodromous, with 5-7 main lateral veins/side; leaves glabrous except for 
scattered hairs on the veins above when young. Inflorescences 4-10(-11) flowered: 
branches glabrous, bearing numerous, caducous, linear, often somewhat curved, mem- 
branous to subherbaceous (green), gland-margined bracteoles; anthesis late March (Gulf 
Coast) to mid-April (Alabama canyon lands). Flowers 15-18 mm diam.; hypanthium ex- 
ternally glabrous; calyx-lobes 3 mm long, narrow-triangular, flaring to a broad base, tip 
somewhat obtuse, margins entire to glandular-denticulate, abaxially glabrous; petals + 
circular, white; stamens 20, anthers pale pink to pale purple; styles 3-4. Fruit 8-10(-12) 
mm diam., subglobose, yellow or flushed pink (occ. reddish); calyx lobes reflexed, on an 
elevated collar; nutlets 3-4, dorsally grooved, sides plane. 

Common name.—Sargent’s hawthorn 

Habitat and Distribution.—Our records are mainly from Alabama, extending west 
to central Mississippi, south to panhandle Florida and eastwards with scattered records 
from Georgia (Fig. 3). It is a locally common species of woodland edges, cut-over wood- 
land and other open woodland. 

Comment.—Crataegus sargentii was the first species to be described in the series and 
this account appears to be the first to characterize it fully. It is most similar to C. gilva but 
that species has paseo leaves and smaller fruit. Occasional more deeply lobed forms 
(LU ca. 15%) may ble oth ies tosome extent, but they will normally retain the 
characteristic overall leaf-form of the species. 


2. Crataegus gilva Beadle. Biltmore Bot. Stud. 1:60. 1902. (Fig. 4). Typr: U.S.A. ALABAMA. Marshall 
Co.: rocky woods, Apr 1901, T.G. Harbison 4374 (LECTOTYPE selected here: A). 
Shrubs or small trees 2-5 m tall; extending shoots glabrous, reddish when young: at 1 yr 
deep reddish-brown, older dark dull gray; thorns fine, 2-4 cm long, straight or slightly 
recurved, dull purplish-brown at 2 yr. Leaves: petioles slender, ca. 20-30% length of blade, 
adaxially grooved, + glandular; blades 3.5-4.5 cm long, broad-lanceolate to narrow-el- 
liptic or very narrow ovate to narrow ovate or narrow ovate-rhombic; tip acute to acumi- 
nate and with 0-3 acute lobes per side these with max. LII usually < 15%: margins sharply 
fine-serrate or occasionally with quite large teeth; venation craspedodromous, lateral 
veins ca. 5-7(-9) per side on average or larger short-shoot leaves; glabrous above and be- 


2) 


— 


BRIT.ORG/SIDA 22 


ar, 
5 


SOONG aig A NS 


980 


TIVAINY 


h\UWY), 


1cm.S. Laurie-Bourque del. 


single leaf at left from extension shoot. Scale bars 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 981 


yee Ee 
ses ecetemeta! 


roe 


& 

| oie eemayeqwes’ 2 
Shed @hancacey 
Her ee eh 


OOS TE 
m4 av eteree py 
SRO 


low except for some hairs on the veins above especially when young. Inflorescences usu- 
ally 3-7 flowered; branches glabrous, densely beset with caducous, narrow-oblong to very 
narrow obovate, membranous to subherbaceous, densely gland-margined bracteoles. 
Flowers (13-)18-20 mm diam. hypanthium externally glabrous; calyx-lobes 6-7 mm long, 
narrow triangular, margins glandular-serrate; petals + circular, white; stamens 20, an- 
thers pale pink; styles 3-4. Fruit ca. 5-8 mm wide at maturity, + orbicular, greenish yel- 
low to yellow; sepals + reflexed; nutlets 3-4. 

Habitat and Distribution.—Crataegus gilva occurs from Mississippi to Georgia and 
northern Florida (Fig. 5), in woodland edges and gaps, clear-cuts and pine-oak woodland. 

Comment.—Crataegus gilva leaves are rather like a sharp-tipped C. mendosa and and 
somewhat resemble a much less clearly lobed C. pinetorum. The leaves are also usually 
somewhat like a proportionately narrower C. sargentii and in the flowering Type they 
are narrow-elliptic, acuminate at the tip, cuneate at the base, virtually devoid of lobes 
and have particularly large marginal teeth, this being the most extreme specimen of the 
species seen. The suspicion that C. gilva may represent an extreme form of C. sargentii is 
heightened by the almost complete lack of known fruiting specimens of the species, the 
fruiting co-type excepted (Harbison 4374/2), which specimen, therefore, does suggest the 
possibility that the leaves often widen after anthesis. 


3. Crataegus mendosa Beadle, Biltmore Bot. Stud. 1:65. 1902. (Fig. 6). Tyre: U.S.A. ALABAMA. Mar- 
shall Co.: near Albertville, 18 Oct 1901, [.G. Harbison 5219 (= 4336/2) (ectotype selected here: US 969431). 


982 BRIT.ORG/SIDA 22(2) 


C. gilva 

C, pinetorum 
Fic. 4 g £1 ry I + Fe a! +. g 9 fT, {¢. | \£ J JTIMAINY ial! ipp In0907 (UWO) Ty pinetorum (bot- 
tom row) from, left to right, Lance 2174 (UWO), Phipps 8173 (UWO) Phipps & 0 8042 (UWO). Scale bars = 1 cm. S. Laurie- 
Bourque del. 


Shrubs or small trees 3-6 m tall; bark of trunk not recorded; l-yr old twigs shiny reddish 
or purplish brown, older dark or dull gray; thorns 2-5 cm long, fine, straight or slightly 
recurved, reddish black at 1 yr, frequent to absent. Leaves: petioles 30-50% length of blades, 
dorsally furrowed, bearing a few small glands; blades +-7 cm long, 2-3 cm wide, broad- 
oblong to elliptic or narrowly ovate; unlobed or with a few obscure lobes; apex subacute 
to blunt on some larger leaves, cuneate at the base; margins with numerous, small, sharp 
teeth; lateral veins 8-9 per side (fewer on smaller leaves), venation craspedodromous; gla- 
brous except for a few hairs on the midvein abaxially when young, dark green. Inflores- 
cences 4-7 flowered; branches glabrous, densely beset with caducous, narrow, semi-her- 
baceous and green to membranous, gland-margined bracteoles. Flowers 14-17 mm wide; 
hypanthium externally glabrous; calyx lobes tr lang ular, abaxially glabrous, margins glan- 
dular-denticulate; petals + circular, white; stamens 20, anthers pink or pink-purple; styles 
3-5. Fruit 8-10 mm diam., + orbicular to broad ellipsoid, yellow-green with pink or salmon 
blush, or red (Beadle); calyx lobes reflexed; nutlets 3-5, dorsally furrowed, laterally smooth. 
Habitat and Distribution.—Scattered from extreme southeastern Texas to west- 

ern Georgia and northern Florida with a disjunct record in South Carolina (Fig. 7), 
Crataegus mendosa is a plant of woodland edges, cut-over woodland and brushy places. 
Comment.—Crataegus mendosa is one of the more distinctive members of the series 
although it is approached in form by C. gilva, which, however, has an acuminate leaf-tip 
and much sharper lobes (if present) and marginal teeth. The extension shoot leaves of C. 
mendosa are ovate to broad-ovate in outline with 3-4 sub-acute to acute lobes that are usu- 
ally quite broad and this can confuse if there are only few typical short-shoot leaves present. 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 983 


Number of Records 
0 


Ty 4 - Crataegus gilva 
LUS Z 


Y a + ! | ee pare ae + rf; +. LS £. eo | Pl 
Fic. 5 y 


3a. Crataegus cf. mendosa Beadle 


Shrubs or small trees 3 m tall; mature bark not recorded; extending twigs glabrous, red- 
dish; at l-yr shiny reddish-brown; older duller dark gray; thorns often on material seen. 
Leaves: petioles ca. 33% length of blade, glabrous, glandular, adaxially grooved; blades 5- 
7 cm long, mostly broad-elliptic in general shape; apex subacute, base cuneate to broad 
cuneate; 0-3 lobed per side, lobes obtuse to subacute, max. LII usually 5-10%; margins 
serrate; venation craspedodromous, ca. 6 main lateral veins per side; both surfaces gla- 
brous. Inflorescences not recorded. Young infructescences with 2-5 fruit, branches gla- 
brous. Young fruit 7-10 mm diam., subglobose to + pyriform, green, glabrous; filament 
bases ca. 20; sepals mainly erose; nutlets ca. 4. 

Habitat and Distribution.—Only known from one locality in Alabama, where it is 
recorded from ‘bottoms’ from where three fine duplicates at DUKE, GA and NO of 
Demaree 50656 (MS: Alabama: Dallas Co., White Oak Creek, PO. Sardis, 29 June 1964) 
show a plant intermediate between C. mendosa, C. opima and C. pulcherrima but which 
cannot be confidently placed with any. More material is needed. 


4. Crataegus pulcherrima Ashe, J. Elisha Mitchell Sci. Soc. 16:77. 1900. (Fig. 8). Typr; US.A. 
FLoripa. Gadsden Co: River Junction, 10-11 Aug 1895, G.V. Nash 2377 (LECOTYPE selected here: DOV 4634; 
ISOLECTOTYPE: US 250142). have selected from Ashe’s co-types the specimens with better foliage and fewer 
fruit rather than more fruit and poorer foliage. 


BRIT.ORG/SIDA 22(2) 


al, 8088, Smith 129 and Lance 2176 (UWO). Scale bars = 1 cm. S. Laurie-Bourque del. 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 985 


FEE Sag ils foes pe 
TE 2 
Spl egy 2 ane 
ae Man ar sn Sait 
eee tae Se 
aaa lgee i at BOA 


?=Crataegus robur Beadle, Biltmore Bot. Stud., 169.1902. Type: U.S.A. FLORIDA. Leon Co.: Tallahasee, 18 Sep 1901, 
C.D. Beadle 2051 (LECTOTYPE selected here: A). 


Shrubs, 2-4 m tall; extending twigs glabrous, green or reddish-tinged; at 1 yr. reddish- 
black; at 2 yr. dark gray; thorns at 2 yr. 2-3.5 cm long, blackish, fine, + straight. Leaves: 
petioles 1-2 cm long, slender, glabrous, glandular, glands sessile or stipitate; blades 2.5- 
5.0 cm long in our material, narrow ovate or + oblong through narrow obovate or nar- 
row rhombic in general shape, with 2-4 shallow obtuse to subacute lobes per side; tips 
acute, bases cuneate; margins serrate with small teeth, somewhat distant for their size; 
venation craspedodromous with 4-5(-6) main lateral veins per side; glabrous above ex- 
cept for along the main veins adaxially young, glabrous below; thin. Inflorescences 4—7- 
flowered with a very short central axis, branches glabrous, bearing plentiful, small, ca- 
ducous, oblong, membranous, stipitate-gland-bordered bracteoles; anthesis mid-April. 
Flowers 15-20 mm diam.; hypanthium externally glabrous; calyx lobes 3-4 mm long, 
narrow-triangular, margins glandular-serrate, abaxially glabrous, tip obtuse; petals + 
circular, white; stamens 20, anthers small, pink; styles ca. 3. Fruit in clusters of 2-5, 6-7 
mm diam., glabrous, deep yellow sometimes flushed red, occasionally purplish-red; ca- 
lyx lobes reflexed, on a short collar; nutlets ca. 3, dorsally grooved, sides plane. 

Common name.—Beautiful hawthorn. 

Habitat and Distribution.—Mississippi to Georgia and northern Florida (Fig. 9). Open, 
rocky woodlands JBP) and swamp borders (Beadle), scarce. 


BRIT.ORG/SIDA 22(2) 


Nitti h /— 
A Se i a ee 
\ tu” J, 7 NY, Ps ape tie 
Mm Na He ae Lt: 
( a % oe if 4 


hipps et al. 8091 (UWO) 


Scale bars = 1 cm. S. Laurie- Bourque del. 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 987 

Vitae por ee om ET Ph EP KL AA OT i S 

HE RR ee ee ee 
gitqeSniomim meer “te ermaueieace esiere 3 
nue BRaSISanay Cee Geecor eee an oa 


%G 
hae che me 
nee 


= 
ai 
Bere 
= 
be 
int 
oy 
= 
st) 
is 


Pilg 
rn 
ib 
Beale 
parat 
ee 
email 
ane 


ae 

& 

nee 
hen 
aa 


Vere 
yA 
ee 

Ss 
ce 
bee 
ee 
i 


fa 41, 1] I dictrihit? £ frat, I-fh i f, ll d qd 
Fic. 9. County p | t 


Comment.—The type species of its series was described by Ashe as having 10 sta- 
mens but neither of Ashe’s co-types (Nash 2377 and Nash 2568) from River Junction, 
Florida confirm this. This species is usually quite easy to distinguish though occasion- 
ally it resembles Crataegus cf. opima. 


5. Crataegus pinetorum Beadle, Bot. Gaz. 28: 343. 1900. (Fig. 4). Type: U.S.A. ALABAMA. De Kalb 
Co.: De Soto State Park, 25 Sep 2001, R. Lance 2174 (NEOTYPE selected here: UWO; ISONEOTYPES: Anniston 
Nat. Hist. Mus., UNC, JSU). 
Shrubs or small trees 2-6 m tall; extending twigs glabrous, with a strong reddish cast; at 
lyr shiny reddish-brown; dull gray-brown at 2 yr; older dull gray; thorns usually present, 
mainly 2-35 cm long, fine, + straight, shiny black or reddish-black at 1 yr, becoming 
black. Leaves: petioles 25-30% of leaf blades, glabrous, sparsely to densely glandular or 
even stipitate-glandular, abaxially grooved; blades 4-6 cm long, nearly full-grown at an- 
thesis, narrowly ovate to elliptic in general shape; tip acute, base cuneate; lobes usually 
2-3 per side, fairly shallow (max. LI] 10-20%), acute; marginal teeth extremely small, acute; 
venation craspedodromous, main lateral veins 5-6 per side; glabrous adaxially and 
abaxially. Inflorescences 2-4-flowered; branches glabrous, bearing plentiful, + persis- 
tent, very narrowly obovate, chartaceous to subherbaceous, densely gland-margined 
bracteoles. Flowers at anthesis not known; hypanthium externally glabrous; calyx lobes 
ca.6 mm long, narrow triangular, margins finely glandular-serrate, abaxially glabrous; 
petals not known; stamens 20, anther color not known; styles porly persistent. Fruit 12- 


988 BRIT.ORG/SIDA 22(2) 


sae 


ee <i ding vw eee 


eye fe eee y 
A) 
Ze ity 


Bale 
ae Bate Ni 


Number of Records 


|__| 0 
Ee] 1-3 


Fic. 10. C | ai 1 . | £ fernt pi t f Ilatod d 


14 mm long, subglobose, dull red; calyx lobes erose on only specimens with mature fruit; 
nutlets 2-4, dorsally grooved, sides plane. 

Habitat and Distribution.—Crataegus pinetorum is found in woodland edges and 
gaps, from south-central Mississippi through Alabama to northern Georgia (Fig. 10). It 
appears to be scarce. 

Comment.—I have been unable to locate Beadle’s original material of this species, for 
which no Type was designated and which were from Marshall Co., Alabama. Nor were 
any specimens labeled ‘pinetorum’ in Beadle’s hand seen so the species, as understood 
here, while it does not disagree with the protologue, could be different from Beadle’s in- 
tention and has therefore received an amplified description here. This is now one of the 
more distinct of the species in ser. Pulcherrimae and, at least superficially, appears most 
similar to C. pulcherrima. It differs from that species in its sharply, rather than obtusely, 
lobed leaves, often densely glandular petioles, exceptionally small leaf marginal teeth 
and red fruit. It also somewhat resembles the more deeply lobed forms of C. gilva. Our 
flowering material was collected after petal drop, consequently a few important charac- 
ters are missing from the description. 

6. Crataegus venusta Beadle, Bot. Gaz. 28:338. 1900. (Figs. 11, 16). Typr: U.S.A. ALABAMA. Jeffer- 
son Co. Birmingham, no date, but probably Sep or Oct 1900, C.D. Beadle 2187/2 (LECTOTYPE selected here: 
US 969379), 20 Apr 1900, C.D. Beadle 2187 (epiryPr selected here: US 969378). 


989 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 


1 cm. S. Laurie-Bourque del. 


be | 


Fic. 11. Li 


8050 (UWO). Scale bars 


990 BRIT.ORG/SIDA 22(2 


?=Crataegus pallens Beadle, Biltmore Bot. Stud. 1:28. 1901. Type: U.S.A. NORTH CAROLINA. Buncombe Co.: 
Biltmore, upland woods, 14 May 1900, Biltmore Herb. BC2 (LECTOTYPE selected here: 9397). 
?=Cratdegus alma Beadle, Biltmore Bot. Stud. 1:64. 1902. Typ: U.S.A. MISSISSIPPL L auderaniec ‘o.: Meridian, no 
date, fruiting specimen; 1G. Harbison H5147 (LECTOTYPE selected here: US 969433). 
Shrubs or small trees +-7 m tall; one-year twigs often deep chestnut; thorns 2-5 cm long, 
usually slender, + straight, very dark at 2-yr old. Leaves: petioles 25-30% length of blade, 
glabrous, very slender, with sparse sessile glands; blades 5-8 cm long at maturity, broad- 
oblong to ovate in general shape; base cuneate to broad-cuneate, apex acute to acumi- 
nate; shallowly lobed, 0-2 lobes per side, LII very shallow to max. ca. 25%, lobes acumi- 
nate young but much blunter in age lobes; margins with numerous, acute to acuminate 
teeth; venation craspedodromous, 5-7 main lateral veins per side; glabrous except for scat- 
tered long hairs adaxially along the mid-vein young. Inflorescences 5-8 flowered; 
branches glabrous, bearing numerous caducous, narrow-oblanceolate to linear, membra- 
nous, clear to greenish, densely gland-bordered bracteoles; anthesis early to mid April. 
Flowers 15-20 mm diam.; hypanthium + glabrous; calyx lobes narrow-triangular, gla- 
brous abaxially, finely gland-bordered; petals + circular, white; stamens 20, anthers pink 
to purplish (or cream, Beadle); styles 3-4. Fruit 8-10 mm diam., suborbicular, greenish- 
yellow to yellow (or reddish, Beadle); calyx lobes spreading or erose; nutlets 3-5, dorsally 
sulcate, laterally smooth. 

Habitat and Distribution.—Texas to Georgia, with one record for Tennessee, the sec- 
ond most northerly record for the series (Fig. 12). It is most abundant in Alabama. Brushy 
forest regrowth, forest gaps and margins. 

Comment.— When flowering, Crataegus venusta is one of the most striking members 
of the series with typical oblong leaf blades, their + lobeless margins bearing numerous 
acuminate teeth. At maturity this characteristic is diminished and typical larger short- 
shoot leaves then have several shallow lobes per side, these + subacute. Crataegus venusta 
was described by Beadle as possessing reddish fruit and cream anthers, but all our mate- 
rial where color determination is possible has yellowish fruit and pink to purple anthers. 

The most distinctive taxon attributed to possible synonymy with C. venusta is C. 
pallens (syn. C.alma),a morphotype with a very sparse and scattered distribution which 
is found from its type locality in western North Carolina to the type locality of its syn- 
onym C. alma in Mississippi. Compared to typical C. venusta it generally has slightly 
smaller leaves (blades typically less than 5 cm long), differently shaped lobes that are 
broader than those of C. venusta with leading edge usually + perpendicular to the midvein 
and less sharp leaf teeth. If C. pallens were to be retained as an accepted species more 
convincing evidence would be required in view of its odd distribution. Nevertheless, in- 
dubitable intermediates with C. venusta are not known. For this reason it is listed sepa- 
rately in the appendix of cited specimens. 


7. ee opune Beadle, Biltmore Bot. Stud. 1:40. 1901. (Fig. 13). Type: US.A. ALABAMA. Butler 
Co. Greenville, 2 Oct 1900, C.D. Beadle 2159/2 (LECTOTYPE selected here: US 981036); C.D. Beadle 2159, 
ae only (EPITYPE selected h 8) 


Crataegus Hide Beadle, eee Bot. Stud. 1:68. 1902. Type: U.S.A. FLORIDA. Leon Co: Tallahasee, 29 Aug. 
1901, 1G. Harbison H4958 = H4059/2 (LECTOTYPE selected here: US 969425), 

?= Pannen contrita Beadle, ae Bot. Stud. Lol. 1902. Type: U.S.A. FLORIDA. Gadsden Co: River Junction, 
3 Apr 1900, C.D. Beadle 2078 (LectToTYPr selected here: US 969333). 

Crataegus illustris Beadle, Biltmore Bot. Stud. 1:68. 1902. Type: U.S.A. Mississippl. Lauderdale Co.: Meridian, - 
Sep 1901, 1G. Harbison H5082 = 4200/2 (LECTOTYPE selected here: US 981041). 


So 
[o) 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 991 


Fic. 12.C ty! I dictvifati € frrt, £ Hatod 4 


Crataegus inanis Beadle, Biltmore Bot. Stud. 1:62. 1902. Type: U.S.A., ALABAMA. Marshall Co: near Albertville, 
May 1901, TG. Harbison H4306 (LECTOTYPE selected here: US 969437). 
?= Crataegus lenis Beadle, Biltmore Bot. Stud. 1:67. 1902. Type: U.S.A., ALABAMA. Butler Co. Greenville, no date, 
fruiting specimen, C.D. Beadle 2153/2 (LECTOTYPE selected here: US 969429). 


Large shrubs or trees to 7 m tall; extending twigs glabrous, reddish-brown, at one year 
blackish-brown, older gray; thorns none to frequent, 2.5-4.0cm long, slender, dark brown 
to blackish at 2-yr old. Leaves: petioles 20-30% length of blade, glabrous, with sparse short- 
stipitate glands; blades 3-5 cm long, mainly broad-ovate to somewhat deltate in general 
shape; base cuneate to broad-cuneate, apex subacute to obtuse; 3-4 main lobes per side, 
the lobes usually + blunt, very shallow to max. LI ca. 20%; margins with small subacute 
teeth; venation craspedodromous, with 4-5 main lateral veins per side: sparsely pilose on 
the mid-vein above when young, glabrous below. Inflorescences 4-7 flowered; branches 
glabrous or sparsely pilose, bearing caducous, narrow-oblong (sometimes curved), mem- 
branous to green, gland-bordered bracteoles; anthesis early to mid April. Flowers 16-18 
mm diam.; hypanthium + glabrous; calyx lobes 4 mm long, triangular, glandular-ser- 
rate at the tip; petals + circular, white; stamens 20, anthers cream to pale or deep purple; 
styles 3-5. Fruit 7-12 mm long, ~ orbicular, yellow or red: calyx lobes erose, or reflexed, 
sitting on a slight collar; nutlets 3-5, dorsally sulcate, laterally smooth. 

Habitat and Distribution.—Extreme southeastern Texas to Georgia, with over half 
the records from Alabama (Fig. 14), this is the most abundant member of the series. It is 
found in open woods and woodland margins. Figure 14 maps C. opima and C. cf. opima 
(sp. 7a) as one. 


BRIT.ORG/SIDA 22(2) 


j 
; LY 4 
fond | 
Koh -| 
ie, /| 
hf: La 


| 


Ma 


| 
aN 


) 


— 
= 


Sr 


——e 


ong 


Dh; eny, 2Nea 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 993 


Number of Records 


Crataegus opima 


f. 1 Nicol dines Lb -t £ fF, + iy ae HY 1 £, Iatad d 
Fic. 14 ty te 


Comment.—The species as defined here comprises all the smaller leaved members of 
the series with broadly proportioned leaves and + blunt lobing. These tend to have few 
and fairly shallow lobes. Both red and yellow-fruited forms are known as are forms with 
cream or purple anthers. It is possible that, as elsewhere in the series, these forms should be 
recognized as distinct taxa, but more detailed study would be required to authenticate this. 


7a. Crataegus cf. opima Beadle (Fig. 15). 


Assigned here are all opima like individuals with ovate to ovate-trullate leaves, 5-7(-8) 
lateral veins per side and 5-8 usually short lobes per side with leaves 5-8 cm long. Forms 
with these larger multiveined leaves may also have acute lobes. More material and fur- 
ther study is required to see whether this should be recognized as a distinct species. 

Note that Figure 14 maps this species and the previous as one and that the list of 
cited specimens (see Appendix) lists only those specimens of this species which are at 
UWO. All these are from Alabama and Missisippi but this entity is undoubtedly much 
more numerous and widespread than this would indicate. 


8. ae tecta Beadle, Biltmore Bot. Stud. 1:26. 1901. (Fig. 16). Type: U.S.A. ALABAMA. Marshall 
ar Albertville, 7 May 1900, T. G. Harbison 2205 (LECTOTYPE selected here: US 969434). The flowering 
era? Harbison 2205at NY displays well the characteristic lobing of the species. However, the fruiting 

co-type at US (TG. Harbison 3285) appears to be C. venusta as does the fruiting specimen at NY. 


?=Crata strina Beadle, Biltmore Bot. Stud. 1:59. 1902. TPE: U.S.A. ALABAMA. Butler Co. Greenville, no 
date, letaae C.D. Beadle 2151 (LECTOTYPE selected here: US 969395) 


BRIT.ORG/SIDA 22(2) 


ae 
Wh AZ 
SEO 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 995 


C. ? venusta 
ey, 
cf. opima 
C. eximia 
Fic 16 1 J g Pa | r I . Sar of +, g ms | £ f. ley rf 3174 (UWO) rs I l.~f¢ #L C ct ! £, DL pp 
ony. onzoanriiany * pu Lnte #1 f 3 £, aie aa ADV STIVAL fr\ - oy ee | wr 4 ee U L 
2205 (US)— middle row, right three; C. eximia from Harbison H4481(A)—bottom row, all. Scale bars = 1 cm. S. Laurie- Bourque del. 


?=Crataegus ancisa Beadle, Biltmore Bot. Stud. 1:63. 1902. Typr: U.S.A. Mississippl Lauderdale Co. Meridian, 16 
Sep 1901, T.G. Harbison H5092 (LECTOTYPE selected here: US 969424). 

Crataegus macilenta Beadle, Biltmore Bot. Stud. 1:64. 1902. TypF: U.S.A. ALABAMA. Marshall Co. Albertville, 7 
Oct 1901, T.G. Harbison H5203= H4283/2 (lectotype selected here: US 969432). 


Shrubs or small trees 4-6 m tall; bark of trunk not recorded; extending shoots glabrous, 
at l-yr dark shiny reddish-black, older dull gray; thorns 2.5-4 cm long, + straight, + fine, 
at 2 yr shiny reddish-black. Leaves: petioles 30-50% length of blade, glabrous, channeled 
above, with a few sessile glands; blades 3-6 cm long, trullate to rhombovate or ovate in 
general shape, some times broadly so; apex subacute, base wide cuneate to cuneate; usu- 
ally fairly deeply (max. LII 20-50%) 3-4 lobed per side, the lobes acute or acuminate to 
somewhat cuspidate, the larger sharper; margins with small acuminate teeth except at 
the base; venation craspedodromous, with 4-5 main lateral veins per side; glabrous above 
except along the main veins when young, glabrous below. Inflorescences 3-6 flowered; 
branches normally glabrous, beset with numerous caducous, sublinear and often wider 
near the apex, sometimes curved, membranous to + herbaceous, gland-margined 
bracteoles. Flowers 17-19 mm wide; hypanthium externally glabrous; calyx lobes ca. 7 
mm long, narrow-triangular, subentire or with margins glandular-denticulate; petals + 
circular, white; stamens 20, anthers pale pink to purple or occasionally ivory to cream 
(see discussion); styles 3-5. Fruit ca.7 mm long, + globose, red or green to yellow at matu- 
rity (see discussion); calyx lobes reflexed; nutlets 3-5, dorsally grooved, laterally smooth. 

Habitat and Distribution.—Crataegus tect Mississippi to western Geor- 
gia, with several records for northern Florida (Fig. 17). It is fairly common in southern 
Alabama and is found in woodland gaps and edges, and brushy places. 


996 BRIT.ORG/SIDA 22(2) 


? v 
= PY, 
= ae 
[ ¢ Tiss 
ri 
\ a ‘ ~ 
MN ‘ 
1. 4 
i ae a 
Y 
o fr 
i 
? ro i 
: a 
pane = oN ; 2 
yw —t 


, Md 

by \ 

HY : ey 
wae 

; Number of Records 

Fic. 17 cr 1 foe L [ff £, late 4 7 


Comment.—The type material of C. tecta is of the form represented by the right hand 
three leaves in the middle row of our figure. This is a very distinctive morphotype in 
series Pulcherrimae. One specimen of C. tecta has a fairly densely pubescent inflores- 
cence. If correctly idetified, this is the only such example for the series. Crataegus ancisa 
and C. macilenta are both recorded by Beadle as having red fruit and pink to purple an- 
thers but do not differ in other respects. Crataegus tecta is most like C. opima but with 
deeper and sharper lobes. Compared to C. incilis the latter species the leaves are propor- 
tionately narrower, have more lateral veins and the anthers not cream or ivory. Some speci- 
mens with deepish lobing, conspicuously acuminate and slightly reflexed lobes are pro- 
visionally referred here rather than to C. venusta. Crataegus opima is the most generally 
similar species to C. tecta, lobe size and shape being the main discriminators. The dubi- 
ous attribution of C. dustrina asa synonym is because the lectotype is quite like 7a, C. sp. 
cf opima and the flowering co-type is rather like C. pallens. In the case of the C. ancisa 
lectotype, this is one of the few unequivocally intermediate specimens encountered, and 
it is between C. opima and C. tecta. 


9. Crataegus incilis Beadle, Biltmore Bot. Stud. 1:41. 1901. (Fig. 18). Type: U.S.A. ALABAMA. Autauga 
Co.: Evergreen, 12 Apr 1900, C.D. Beadle 2143 (LECTOTYPE selected here: US 969493). 


Crataegus concinna Beadle, Biltmore Bot. Stud. 1:70.1902. TyPF: U.S.A. FLORIDA. Li 


perty Co. Bristol, 2 Apr 1901, 


‘G. Harbison H4040 (LECTOTYPE selected here: US 981039). 


KY 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 997 


Fic, 18. Line drawing of C. incilis from flowering specimen Phipps 7891 (UWO), fruiting speci Phipp 
Phipps 7844 (UWO). Scale bars = 1 cm.R. Bremmer del. 


Shrubs or small trees 4-7 m tall; l-yr old twigs dark red brown, at 2 yr dark gray; thorns 
1.5-3cm long. Leaves: petioles 1.5-2.5 cm long, slender, glabrous, with few to several sessile 
glands; blades 3.5-5.5(-7.5) cm long, trullate to narrow-ovate in general outline; subacute 
at tip, base cuneate, then tapering into the upper winged part of the petiole; 3-4 lobes per 
side with max. LIT 15-40%, lobes usually + narrow-c uspidate; margins with small some- 


998 BRIT.ORG/SIDA 22(2) 


what distant subacute teeth; venation craspedodromous, with about 5-7 pairs of lateral 
veins/side; glabrous on both sides at maturity. Inflorescences 3-7 flowered; branches gla- 
brous; bracteoles abundant, caducous, linear. I ,gland-margined. Flowers 14- 
18 mm diam.; hypanthium glabrous; calyx lobes ca. 7 mm long, narrow triangular, mar- 
gins subentire to glandular-denticulate; petals white, + circular; stamens 20, anthers pink; 
styles 3-5. Fruit 5-8(-10) mm diam.,, globose, red or yellow; calyx lobes wide-spreading; 
nutlets 3-5, dorsally grooved, laterally smooth. 

Habitat and Distribution.—Most of our records are from northeastern Alabama, but 
it is known from Mississippi to southwestern Georgia and northwestern Florida (Fig. 19). 
Crataegus incilis is found in open woods and woodland margins. 

Comment.—The deeply cut trullate leaves, with the acute tips of their triangular lobes 
ina + straight line in the more extreme forms, as in our illustration, are very distinctive. 
The Type, and most specimens seen, however, do not have quit nearly straight-side 
lobes as seen in the illustration and their lobes may be described as + narrow cuspidate. 
Forms with somewhat more rounded lobes and yellow fruit are perhaps represented by 
C. concinna. However, it should be noted that the fruiting co-types of C. concinna at US 
(both H4040/2) appear to be a different species, one with barely lobed leaves. The deli- 
cate leaves and twigs of this species are peculiarly attractive. 


10. Crataegus eximia Beadle, Biltmore Bot. Stud. 1:62. 1902. (Fig. 16). Type: U.S.A. ALABAMA. 

Jackson Co. Sand Mountain, near Pisgah, 6 May 1900, T.G. Harbison 4448 (LECTOTYPE selected here: A). 
Shrubs or small trees 3-4 m tall; l-year old twigs reddish-brown; older gray; thorns 3-4 
cm long, usually fine, slightly curved, at 2 yrs. blackish-gray. Leaves: petiole 25-40% length 
of blade, glabrous, adaxially grooved, densely beset with stipitate glands, at lest young; 
blades mainly 4.5-6.5 cm long at anthesis, to 8 cm at maturity, ovate-rhombic to ovate- 
trullate; 3-4(-5) lobes per side, these subacute to acute, max LI] usually 20-33%; margins 
finely dentate; venation craspedodromous, 5-6 main lateral veins per side; abaxial sur- 
face scabrous-hairy only along main veins when young, glabrous below. Inflorescences 
4-6 flowered; branches glabrous, bearing caducous, very narrowly obovate, + membra- 
nous, gland-margined bracteoles. Flowers 22-25 mm diam.; hypanthium glabrous; calyx 
lobes 4-5 mm long, finely glandular-serrate, glabrous on both sides; petals + circular, 
white; stamens ca. 20, anthers cream; styles 3-4. Fruit (Beadle) 8-12 mm wide, globose, at 
maturity reddish; nutlets 3-5 (Beadle), dorsally grooved. 

Habitat and Distribution.—Rocky woods and glades, Sand Mountain, Jackson Co., 
Alabama (Fig. 20). 

Comment.—Known to me JBP) only froma flowering co-type, this nevertheless seems 
one of the most distinct entities in the series on the basis of its large rhombovate leave 
with deep and angular lobing, large flowers and late to very late flowering date (23 May 
in 1901). Series Pulcherrimae are generally scarce this far north. 


APPENDIX 

Further cited specimens (species, states and counties arrangement alphabetic). 
percep gilva Beadle 

ALABA Cherokee Co.: Little R.Canyon North Park, 24 Apr 1969, KE. Landen & J. Cofields.n. JSU); Little River 
Canyon eee ca. 1 km direct NW from Daniel's Gap, 34° 18'N,85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 
7937a (UWO); entrance road to Little River Canyon Mouth Park, 33° 17'N, 88° 40.5'W, 18 Apr 1999,J.B. Phipps & RJ. 
O’Kennon 7922 (UWO); entrance road to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 1999, J.B. 
Phipps & RJ. O’Kennon 7923 (UWO); Little River Canyon rim road ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 999 


aw 


Number of Records 
0 


Crataegus incilis 


Fic. 19 fa 4,1 LAictrihnt) £ on tlie £, Hatod dq 
7 r J 


85° 42'W, 22 Sep 1999, J.B. Phipps & RJ. O’Kennon 8046 (UWO); Little River Canyon rimroad ca. 1 km direct NW 
from Daniel’s Gap, 34° 18'N, 85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7940 (UWO). De Kalb Co.: Along 
Little R.in Little R.Canyon, near Eberhart’s Point; 20 Apr 1968, K.E. Landen & J. Cofield s.n. (JSU); De Soto State Park, 
campground area, 34° 30'N, 85° 38'W, 19 Apr 1999, J.B. Phipps & RJ. O’Kennon 7956 (UWO); De Soto State Park, 
campground area, 34° 30'N, 85° 38'W, 19 Apr 1999, .B. Phipps & RJ. O’Kennon 7954 (UWO). Marshall Co.: no 
location, no date, C.D. Beadle s.n. (A). Tallapoosa Co.: Along Co. Road 2,0n hilltop S of Romulus, scrubland on W 
side of road, 10 Apr 1999,R. Lance s.n. (UWO); Co.Rd. 2,10 km NE of jct.10 or 51,SE side at gate with big overview 
opposite, 33° 13'N, 87° 42'W, 13 Apr 1999, /.B. Phipps 7889 (UWO). Tuscaloosa Co.: near Northport Econolodge, 
wooded bank behind motel, at edge, 33° 14'N,87° 34'W, 11 Apr 2000, J.B Phipps 8118 (UWO). 

FLORIDA. Gadsden Co.: Aspalaga, prety R.,05 Apr 1964, 5. McDaniel 4095 (IBE); Greensboro, 05 
Apr 1964, D. Demaree 49482 (NO). Jackson Co.: Three Rivers State Park, road leading to camping area no.1,1.m 
N of Sneads, 03 June1983, R.K. Godfrey 80684 (UWO). Leon Co.: near Tallahassee, Apr 1843, Rugel 44 (BM, vs 
mesic woodland on bluff by Lake Miccosukee, Ring Oak Plantation, E of Miccosukee, 22 Mar 1983, R.K. Godfrey 
80354 (UWO). Liberty Co.: Bristol, Appalachicola R., 24 Mar 1967, S. McDaniel 8625 (IBE). 

GEORGIA. Burke Co.:Rte 56,N of intersection with 80,N of Lake Crystal Rd. to E,33° 05'N, 82° 05'W,09 Apr 
1991,J/.B. Phipps 6498 (UWO). Clarke Co.: Athens, Ag. Campus, Oconee Hillside, 11 Apr 1929, .R. D E802 
(GA, 2). Houston Co.: Oakey Woods Wildlife Management Area, just S of Kathleen, E side of gravel road to check- 
in station, 0.4 mi S of fork, 32° 28.680'N, 83° 34.667'W, 10 Apr 2001, R. Lance 2130 (UWO).Merriwether Co.: White 
Sulphur Springs, 26 May 1924, 7.G. Harbison 16307 (UNC). Rabun Co.: Tallulah River State Park, near overlook to 
dam, NE of dam,N 34° 44.429", W 83° 23.655’, 26 Apr 2001,R. Lance 2135 (UWO). Randolph Co.: SE of Coleman, ca. 
0.5 mi W of Co. Rd. 160, on N side of Co. Rd. 62, across from old drive, 31° 28.630'N, 84° 52.875'W,09 Apr 2001,R. 
Lance 2112 (UWO); SE of Coleman,0.5 mi W of jct.with Hwy 160 on S side of road, at woods and field corner, near 
phone box"616-6,"31° 39.898'N, 84° 53.642'W, 10 Apr 2001,R. Lance 2120 (WO). Richmond Co.: Augusta, sand 
hills, 20 Apr 1900, S.7. Olney and J. Metcalf 222 (DOV); Augusta, no date, A. Cuthbert 2 (GH). 

MISSISSIPPI. Greene Co.: roadside in wooded country near McLain on US 98, 31° 07'N, 88° 48'W, 07 Apr 


— 


‘ 


1000 BRIT.ORG/SIDA 22(2) 
Serie 


GPRM CGD. aneeee a cereaia cae 
Seer naee cel 


rion 4 K 
ye nan erie 
ae ey: mate es eens 
pte 


et 
ae 
=o 


E 


ae | 
SeeaNe eee ey 
aries, ay 


(1 
a 
ame 
we 


a 


Hoe 
et co 
irene fat 


Number of Records 


0 
fee] 1-3 


Fic 70. yl i 4 th Ly Poe aes eee Ilatad d 


1984, J.B. Phipps &T.C. Wells 5335 (UWO); roadside in wooded country near McLain on US 98, 31° 07'N, 88° 48'W, 
07 Apr 1984, J.B. Phipps & T.C. Wells 5339 (UWO). Jones Co.: Laurel, just N of jct. Tallahoma/Tallahala Creeks, NW4 
535, 02 Apr 1980, 5. McDaniel s.n. (UNA, UWO). Lauderdale Co.: Meridian, 12 Apr 1900, CS. Sargent s.n. (DOV). 
Simpson Co.: Saratoga, 01 Apr 1903, S.M. Tracy 8490 (MO). 


Crataegus incilis Beadle 
ALABAMA. Calhoun Co.: Brown Ridge Road, 0.3 mi SW of jct. with county highway 55, scrubby hill E side of 
Brown Ridge Road, opposite open field, 08 Apr 1998, R. Lance s.n. (UWO). Cherokee Co.: Little River Canyon 
rimroad,ca. 1 mi NW of Daniel’s Gap, 34° 17'N,85° 41'W,08 Oct 1998, J.B. Phipps 7844a (UWO); Little River Canyon 
rimroad,ca. 1 mi NW of Daniel's Gap, 34° 17'N, 85° 41'W,08 Oct 1998, J.B. Phipps 7844c (UWO); Little River Canyon 
rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7939 
(UWO); Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 22 1999, J.B. Phipps 
& RJ. O’Kennon 8040 (UWO); entrance road to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 1999, 
J.B. Phipps & RJ. O’Kennon 7946 (UWO); Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34° 
18'N, 85° 42'W, 22 Sep 1999, J.B. Phipps & RJ. O’Kennon 8048 (UWO); Little River Canyon rimroad ca. 1 km direct 
NW from Daniel’s Gap, 34° 18'N, 85° 42'W, 22 Apr 1999, J.B. Phipps & RJ. O’Kennon 8039 (UWO); Little River Canyon 
rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’kennon 7942 
(UWO); Little River Canyon rimroad ca. 1 km direct NW from Daniel’s Gap, 34° 18'N, 85° 42'W, 22 Sep 1999, /.B. 
Phipps & R.J.O’Kennon 8045 (UWO); Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 
85° 42'W, 18 Apr 1999, J.B. Phipps & R.J.O’Kennon 7944 (UWO); entrance road to Little River Canyon Mouth Park, 
34° 17'N, 88° 40.5'W, 18 Apr 1999, 1B. Phipps & RJ. O’Kennon 7924 (UWO); Little River Canyon rimroad ca. 1 km 
NW from Daniel's Gap, 34° 18'N, 85° 42'W, 22 Sep 1999, /.B. Phipps & RJ, O’Kennon 8042 (UWO). Dallas Co.: 
road to Elm Bluff from Shephardville, ca. 1.5 km W of Shephardville, 32° 11'N,87° 05'W, 24 Apr 1999, R.J.O’Kennon 
&R. Lance 14458 (UWO); Elm Bluff, near Alabama River, 32° 11'N,87° 06'W, 02 Oct 1998, J.B. Phipps & R. Lance 7795 
(UWO). Monroe Co.: Haines Island, high limestone ridges along Alabama R., Red Hills area, 05 Apr 1986, A.R. 


es 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 1001 


Diamond 2256 (AUA). Tuscaloosa Co.: Lake Lurleen State Park ded slope near campground, 10 Apr 1999,R. 
Lance 66 (UWQ) 

FLORIDA. Gadsden Co.: Hardin Heights, jct. FL 269 and E extension of Brickyard Rd.,03 Aug 1983, J.B. 
Nelson & G.R. Knight 2449 (FSU). Liberty Co.: Bristol, 31 Mar 1902, 1.G. Harbison H4040 (A). Washington Co.: FL 
273, a mi ue of Chipley, 14 Apr 1966, J. Beckner & W. D’Arcy 1160 (DUKE). 

ORGIA. Mcintosh Co.: Meridian, 19 Apr 1912, 1.-G. Harbison 10859 (NCU). 
Pre ss Co.: SE of Coleman, ca.0.5 mi W Co. Rd. 160, on N side of Co. Rd. 62, across from old drive, N ais 
28.630', W 84° 52.875',09 Apr 2001,R. Lance 21 abies: SE of Saleen ca.6.7 mi from S end of Rd. 20 and 1.1 
ae with Hwy 160,ca.20 ft W of road, in dland, just W of cemetery and telephone cable post 
”N 31° 39.708' W 84° 54.131', 09 Apr 2001, R.Lance 211 9 (UWO). 

Beane Smith Co.: SE of Lemon on 35, SR 533A, 32° 07'N, 89° 29'W, 26 Sep 1999, 1B. Phipps, RJ. 

O’Kennon & R. Lance 8090 (UWO) 


Crataegus mendosa Beadle 
ALABAMA. Cherokee Co.: Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34°18'N 85° 42'W, 
18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7941 (UWO);Little River Canyon rimroad ca.1 km direct NW from Daniel's 
Gap, 34° 18'N 85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7938 (UWO); Little River Canyon Park, just N of 
Powell Trail access on rim road, 34° 19.729'N 85° 40.232'W, 25 Sep 2001,A.Lance 2176 (UWO); Little River Canyon 
rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N 85° 42'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7939a 
(UWO). Choctaw Co.: AL 10,9 km SE of jct.Co. Rd. 9 (at Lisman), SW side of road, 32° 06'N 88° 15'W, 14 Apr 1999, 
J.B. Phipps 7904 (UWO). Dallas Co.: road to Elm Bluff from Shephardville, ca. 1.5 km W of Shephardville, 32° 11'N 
87° 05'W, 24 Sep 1999, RJ. O’Kennon & R. Lance 14455 (UWO); road to Elm Bluff from Shephardville, ca. 1.5 km W 
of Shephardville, 32° 11'N 87° 05'W, 24 Sep 1999, RI. O’Kennon &R. Lance 14454 (UWO). Madison Co.: Pine Bluff 
(1315 ft), 10 Apr 1882, C. Mohr 446 (ALU). Pickens Co.: On dirt road joining Fellowship and New Hope churches, 
2.2 mi N of Co.Road 30, NW 1/4 Sec 24T19S R14W,09 Apr 1977,L.J. Davenport 22 (UNA). Tallapoosa Co.: Approx 
15 mi from Dadeville post office, SSW, 4 mi from Martin Dam, 01 May 1971, EJ. Widder 096 (AUA); Route 50 ca. 1 
km E of Martin Dam, 32° 41'N 85° 53'W, 16 Oct 1984, J.B. Phipps 5500 (UWO). Tuscaloosa Co.: Co. Rd. 2,5 km NE 
of jct.10 or 51,33° 12'N 87° 43'W, 13 Apr 1999, J.B. Phipps 7886 (UWO);Co.Rd.2 at 11 km NE jct.51 at overlook, 33° 
13'N 87° 42'W, 23 Sep 1999, J.B. Phipps & RJ. O’Kennon 8068 (UWO); Co .Rd. 2 at 11 km NE jct.51 at overlook, 33° 
13'N 87° 42'W, 23 Sep 1999, .B. Phipps & RJ. O’Kennon 8070 (UWO); Co.Rd. 2 at 11 km NE jct.51 at overlook, 33° 
13'N 87° 42'W, 23 Sep 1999, .B Phipps & RJ. O’Kennon 8065 (UWO):Co.Rd. 2,10 km NE of jct.10 or 51,33° 13'N 87° 
42'W, 13 Apr 1999, J.B. Phipps 7892 (UWO); Kellytown Rd., 1 km from jct. Valley Rd., 33° 07'N 87° 49'W, 23 Sep 1999, 
J.B. Phipps & RJ. O'Kennon 8073 (UW 

FLORIDA. Gadsden Co.: River ee 06 Sep 1940, WA. Murrill 20 (MO). Jackson Co.: ca.9 mi N of Marianna, 
21 Oct 1955, R.K. Godfrey & H. Kurz 54277 (NCSC). Leon Co.: 4 mi E of Tallahassee (decid. woods), 16 Mar 1955, R.K. 
Godfrey 53022 (USF).Wakulla Co.: near Wakulla Springs, 12 Apr 1931, EJ. Palmer 38571 (MO). 

ORGIA. Randolph Co.: SE of Coleman, ca.0.5 mi W of Co. Rd 160,0n N side of Co. Rd.62 on margin of 

private drive, N 31° 28.630', W 84° 52.875', 27 Sep 2001,R. Lance 2194 (UWO); SE of Coleman, W side of Co. Rd 20 
(Cemetery Road), ca.6 mi from its S end and 1.8 mi from its jct. with Hwy 160, across from pine plantation and 
“posted” sign, N 31° 39.375', W84° 54.853', 27 Sep 2001.R. Lance 2182 (UWO); SE of Coleman, 6.2 mi from S end of 
Rd. 20 and 1.6 mi from jct. with Hwy 160, N 31° 39.468", W 84° 54.642', 27 Sep 2001,R. Lance 2183 (UWO). 

LOUSIANA. Rapides Parish: Lacompte Quad.,09 Jul 1977, L.E. Urbatsch 3237 (LSU). St. Tammany Parish: 
4 mi W of Bush, N of La 40, pinewoods, 26 Sep 1975, R. D. Thomas & C. Allen 47164 (NLU). Washington Parish: 
Bogalusa, 03 Oct 1919,R.S. Cocks 3157 (NO); woods at Mt. Hermon Cemetery beside La.450 NW of Mt. Hermon, 
Sec. 40,115, R9E, 14 Jun 1983, R.D. Thomas 84147 (UWO) 

eres Covington Co.: 2.5 mi NE of Collins, pine hardwoods, 08 Apr 1961,L.H. Shinners 29359 (GA). 
George Co.: Pascagoula river S of MS 26,2 mi E.of Benndale, 31° 07'N, 88° 48'W, 28 Mar 1975, R.D. Thomas, CM 
Allen & G. as 42999 (LSU). Greene Co.: roadside in wooded country near McLain on US 98, 31° 07'N, 88° 
48'W,07 Apr 1984, J.B. Phipps & T.C. Wells 5338 (UWO); roadside in wooded country near McLain on US 98,07 Apr 
1984, J.B. Phipps & T.C. Wells 5336 (UWO). Kemper Co.: 2 mi E. of De Kalb, 12 Aug 1961, S. McDaniel 2669 (\BE). 
Lauderdale Co.: Meridian; 12 Apr 1900, C.S. Sargent s.n. (DOV); 3 mi S of Meridian, 19 Jul 1956, .D. Ray, Jr. 7087; 
(USF). Noxubee Co.: Butler Rd., running N from MS 21 ca.1 miW of Gholson, 32° 57'N, 88° 45'W, 27 Sep 1999, J.B. 
Phipps, R.J. O’Kennon & R. Lance 8101 (UWO); turn N from Hwy 21 at Gholson onto gravel road beside Church of 
God of Prophecy, collections along road, 1.5 mi N of Hwy 21,T13NR15E $26, 27 Apr 1984, 7.£. Smith 273 (UWO). 
Perry Co.: damp wooded roadsides on US 98 near New Augusta, 31° 13'N, 89° 00'W, 07 Apr 1984, J.B. Phipps & 
T.C. Wells 5342 (UWO). Simpson Co.: Saratoga, 01 Apr 1903, 5.M. Tracy 8490 (MSC); SE of Lemon on 35, SR 533A, 
32° O7'N, 89° 29'W, 26 Sep 1999, J.B Phipps, RJ. O’Kennon & R. Lance 8088 (UWO). Winston Co.: ca. 8.0 mi SE of 


— 


1002 BRIT.ORG/SIDA 22(2) 


Louisville on sides of NW-SE gravel rd.ca. 1 mi SE of int.with N-S FAS blacktop, T14N R14E 533 SW4,08 May 1985, 
TE. Smith 1291 (OWO);ca. 2.5 mi ENE of Handle, side of NE-SW gravel rd.ca.0.1 mi NE of jct. with NW-SE gravel rd., 
T13N R14E S14 SE4, 24 Jun 1985, LE. Smith 1403 (UWO). 
SOUTH CAROLINA. Newberry Co.: 21 Apr 1971, 1.H. Buff Jr 203 (USCH). 

S. Jasper Co.: Rich wooded slopes, road to Magnolia Springs,8 mi S of Jasper, 11 May 1945,R.McVaugh 
6833 (BRIT). Newton Co.: Saw Mill Road, N side, 30° 52'N, 93° 50'W, 18 Oct 1988, J.B. Phipps 6273 (UWQO): wooded 
hills, old pine-oak stand, 4 mi W of Newton, 13 May 1945, R. McVaugh 6855 (BRIT); wooded hills, old pine-oak 
stand, 4 mi W of Newton, 13 May 1945, 8. McVaugh 6857 (BRIT). 


Crataegus opima Beadle 
A MA. Autauga Co.: Hwy 45,6 miE of Prattsville, 21 Apr 1962, 7. Denton 81 (ALA). Butler Co.: near Greensville, 
16 Apr 1931, EJ. Palmer 38688 (MO). Cherokee Co.: Little River Canyon rimroad ca. 1 km direct NW from Da Le 
Gap, 34° 18'N, 85° 42'W, 22 Sep 1999,J.B. Phipps & RJ. O’Kennon 8043 (UWO), Little River Canyon rimroad ca. 
direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 22 Sep 1999, J.B. Phipps & RJ. O’Kennon 8047 (UWO); Little aa 
Canyon Mouth Park, entrance Road, 34° 17'N, 85° ee W, 22 Sep 1999, /.B. Phipps & RJ. O’Kennon 8056 (UWO); 
entrance to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7930 
(UWO); Little River Canyon rim road ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 18 Apr 1999, /.B. 
Phipps & R.J.O’Kennon 7937 (UWO); entrance road to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 
1999, J.B. Phipps & RJ. O’Kennon 7926 (UWO); Little River Canyon one third way up access road to top, beyond 
Canyon Mouth Park, 34° 17'N, 85° 41'W, 08 Oct 1998, J.B. Phipps 7841 (UWO); Little River Canyon one third way 
up access road to top, beyond Canyon Mouth Park, 34° 17'N, 85° 41'W, 08 Oct 1998, J.B. Phipps 7840 (UWO). 
Geneva Co.: Bluff on E side of Hwy 45 at crossing of Chectawhatchee River, near Bellwood, 23 Sep 1999, 8. Lance 
s.n. (UWO). Conecuh Co.: US 31 at Travis Bridge, Sepula R., 17 Aug 1985,A.R. Diamond 1517 (AUA). Dallas Co.: 2 
mi NW Shephardsville, SE bank Alabama R., 08 Sep 1982, 8.C.Gunn 1231 (AUA); Elm Bluff, W of Shephardville on 
Alabama River, 32° 11'N, 87° 05'W, 02 Oct 1998, J.B. Phipps & R. Lance 7791 (UWO); road to Elm Bluff from 
Shephardville, ca. 1.5 km W of Shephardville, 32° 11'N, 87° 05'W, 24 Sep 1999, RJ. O’Kennon & R. Lance 14456 
(UWO); road to Elm Bluff from Shephardville, ca. 1.5 kn W of Shephardville, 32° 11'N, 87° 05'W, 24 Sep 1999 RJ. 
O’Kennon & R. Lance 14457 (UWO); road to Elm Bluff from Shephardville, ca. 1.5 km W of Shephardville, 32° 11'N, 
87° 05'W, 24 Sep 1999, RJ. O’Kennon & R. Lance 14459 (UWO); road to Elm Bluff from Shephardville, ca. 1.5 km W 
of Shephardville, 32° 11'N, 87° O5'W, 24 Sep 1999, RJ. O’Kennon & R. Lance 14461 (UWO): road to Elm Bluff from 
Shephardville, ca. 1.5 km W of Shephardville, 32° 11'N, 87° 05'W, 24 Sep 1999, RJ. O’Kennon & R. Lance 14460 
(UWOQ); Elm Bluff,W of Shephardville on Alabama River, 32° 11'N,87° 05'W,02 Oct 1998,/.B. Phipps & R. Lance 7792 
(UWQ); Elm Bluff, W of Shephardville on Alabama River, 32° 11'N,87° 05'W,02 Oct 1998,J.B. Phipps &R. Lance 7793 
(UWO); Dallas Co.Rd.407 between Shephardville and Elm Bluff near parking area on right, along N-W trending 
track at 1 km, 32° 11'N,87° 05'W, 14 Apr 1999, J.B. Phipps 7903 (UWO). Monroe Co.: near Frisco City;01 Jun 1940, 
J. Kelly 22 (GA). Tallapoosa Co.: Route 50 ca. 1 km E of Martin Dam, 32° 41'N, 85° 53'W, 16 Oct 1984, /.B. Phipps 
5497a (UWO). Tuscaloosa Co.: Tuscaloosa, 11 May 1890, C. Mohr s.n. (ALU); Romulus Rd. (Co. Rd. 51) at 3.3 km 
short of Gainesville Rd, 33° 08'N, 86° 44'W, 23 Sep 1999, J.B. Phipps & RJ. O’Kennon 8057 (UWO); Romulus Rd. (Co. 
Rd. 51) at 3.3 km short of Gainesville Rd., 33° 08'N, 86° 44'W, 23 Sep | 999, J.B. Phipps & RJ. O’Kennon 8058 (UWO); 
Co.Rd. 2,10 km NE of jct.10 or 51, SE side of, at gate with big overview opposite, 33° 13'N,87° 42'W, 13 Apr 1999, 
J.B. Phipps 7890 (UWO); Co. Rd. 2,ca.3 mi S of Coker, E side of road on outside of curve, 33° 13.5'N,87° 46'W, 13 Apr 
1999,J.B. Phipps 7893 (UWO); Co.Rd. 2,3 km NE of jct.10 or 51, SE side of road just before es house, 33° 10'N, 87° 
43'W, 13 Apr 1999,1.8. Phipps 7885 (UWO); Kellytown Rd., 1 km from jct.Valley Rd., 33° 07'N, 87° 49'W, 23 Sep 1999, /.B. 
Phipps & RJ. O’Kennon 8075 (UWO). Sumter Co.: ca.9 mi SSE of York, by Ala 17,05 Jun 1971,R. Kral 42979 (VDB). 
FLORIDA. Gadsden Co.: Off Dolan Rd. to N, WSW of River Jct., 30 Apr 1983, R.K. Godfrey 80540 (UWO). 
GEORGIA. De Kalb Co.: Decatur, 10 Aug 1901, R.M. Harper 1184 (MO). Early Co.: Kolomoki Mounds State 
Park, oak woods just W of museum, 31° 28'N, 84° 55'W, 03 Oct 2000, J.B. Phipps & R. Lance 8196 (UWO): Kolomoki 
Mounds State Park, oak woods just W of museum, 31° 28'N, 84° 55'W, 03 Oct 2000, J.B. Phipps & R. Lance 8195 
(UWO). Grady Co.: Susina Plantation, 11 Apr 1966, R. White 203 (IBE). Randolph Co.: SE of Coleman, 6.7 mi from 
S end of Co. Rd. 20 and 1.1 mi from jct. with Hwy 160, ca. 20 ft.W of road, in young oak woodland just W of 
cemetery and telephone cable post"616-12,"N31° 39.708", W 84° 54.131', 27 Sep 2001,R. Lance 2185 (UWO); Co. 
Rd. 160, just S of Coleman, 31° 39.5'N, 84° 53'W, 03 Oct 2000,/.B. Phipps & R. Lance 8198 (UWC); SE of Coleman, ca. 
0.5 mi W of Co.Rd 160,0n N side of Co.Rd.62, across from old drive, N 31° 28.630',W 84° 52.875', 27 Sep 2001,R. 
Lance 2188 (UOWQ);SE of Coleman, ca.0.5 mi W of Co.Rd 160, 0n N side of Co. Rd. 62, across from old drive, N 31° 
28.630',W 84° 52.875',09 Apr 2001, R. Lance 2109 (UWO). 
LOUISIANA. Washington Parish: Thigpel Cr, N of LA 16 and E of Enon, 09 Apr 1978, R.D. Thomas, D. More- 
land & P Pias 57458 (NLU). 


A 


— 


eae 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 1003 


MISSISSIPPI. Clarke Co.: Clarko State Park, wooded slope near exit drive on US Hwy 45,09 1999, R. Lance 
s.n. (UWO); Clarke County State ee N exit on US 45, ol 42'N, 88° 07'W, 12 Apr 1998, J.B. Phipps 7698; (UWO). 
meet! Co.: Meridian, 27 Apr 1901, IG. Harbison H4200 (US).Noxubee Co.: Butler Rd.,running N from MS 21 

.1 miW of Gholson, just after first te road left, 32° 57" N,88° 45'W, 16 Apr 1999, J.B. Phipps 7915 (UWO); Butler 
4 ee N from MS 21 ca.1 mi W of Gholson, E side of road, 32° 57'N, 88° 45'W, 27 Sep 1999, J.B. Phipps, R.J. 
O’Kennon &R. Lance 8102 (UWO); Butler Rd., running N from MS 21 ca. 1 miW of Gholson,E side of road, 32° 57'N, 
88° 45'W, 27 Sep 1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8096 (UWO); Butler Rd., running N from MS 21 ca.1m 
W of Gholson, E side of road, 32° 57'N, 88° 45'W, 27 Sep 1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8099 (UWO); 
Butler Rd. running N from MS 21 ca.1 miW of Gholson,€ side of road, 32° 57'N,88° 45'W, 27 Sep 1999, J.B. Phipps, 
RJ. O’Kennon & R. Lance 8098 (UWO); Butler Rd., running N from MS 21 ca.1 mi W of Gholson, just after first side 
road left, at right, 32° 57'N, 88° 45'W, 16 Apr 1999, J.B. Phipps 7916 (UWO). Perry Co.: ca. 11 mi NE of Wiggins, NW 
of jct. of forest road 318B-1 and Black Creek Trail, along Black Creek Trail,"Red Hills", 11S, R10W, section 28 and 21, 
30 Apr 19??, REC. Nacziet al 5119 (VDB). Smith Co.: SE of Lemon on 35,SR 533A,32° 07'N,89° 29'W, 26 Sep 1999, 
J.B. Phipps, R.J.O’Kennon & R. Lance 8089 (UWO). Winston Co.: ca. 12 mi ESE of Louisville N side of SE-SW gravel rd. 
ca. 1 mi NW of Hwy 490 int, T14N R14E $23 SE4,08 May 1985, TE. Smith 1289 (UWO). 

TEXAS. Jasper Co.: S of Jasper, 15 Apr 1942, C.L.& AA. Lundell 11191 (SMU); S of Jasper off highway 96 
between road to Magnolia Springs and road to Roganville, E side of road, 14 Apr 1963, D.S.and H.B. Correll 27231 
(LL); Road to Magnolia Springs, 8 mi S of Jasper, 11 May 1945, R.McVaugh 6833 (SMU); S of Jasper; 09 Sep 1942, 
CL. Lundell & S.W. Geiser 1117819 (SMU). Newton Co.: Newton, 04 Jul 1960, /. Knapp s.n. (NATC); Saw Mill Road ca. 
1.5 km W of Saw Mill Town, 30° 52'N, 93° 50'W, 12 Apr 1988, J.B. Phipps & L. Lowrie 6079 (UWO); Saw Mill Road ca. 
1.5 km W of Saw Mill Town, 30° 52'N, 93° 50'W, 12 Apr 1988,J.B. Phipps & L. Lowrie 6077 (UWO); Saw Mill Road, N 
side, 30° 52'N 93° 50'W; 18 Oct 1988, J.B. Phipps 6274 (UWO); Saw Mill Road, N side, 30° 52'N, 93° 50'W; 18 Oct 
1988, J.B. Phipps 6275 (UWO); Saw Mill Rd.ca.0.75 km W of Saw Mill Town, 30° 52'N, 93° 49'W, early Nov 1987, L. 
Lowrie s.n. (UWO); intersection of two dirt roads S of US 190, the one joining US 190 3 mi W of Newton, 30° 52'N, 
93° 51'W, 12 Apr 1988, J.B. Phipps & L. Lowrie 6076 (UWO); Saw Mill Road, N side, 30° 52'N, 93° 50'W, 18 Oct 1988, 
J.B. Phipps 6272 (UWO); Saw Mill Road, N side, 30° 52'N, 93° 50'W, 18 Oct 1988, J.B. Phipps 6276 (UWO); 4 mi W of 
Newton, 13 May 1945, 8. McVaugh 6856 (SMU). 


Crataegus cf. opima Beadle 
ALABAMA. Cherokee Co.: Entrance road to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 1999, 
J.B. Phipps & RJ. O’Kennon 7945 (UWO); entrance road to Little River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 
Apr 1999, J.B. Phipps & RJ. O’Kennon 7927 (UWO); Little ie Canyon, S rim above Canyon Mountain Park, be- 
tween mi 2 and 3 at county road bord nrim,07 Apr 1998, R. Lance s.n. (UWO). Dallas Co.: r oad to Elm 
Bluff from Shephardville, ca. 1.5 km W of snegharcvile along trail to deer i (N of road), 32° 11'N, 87° 05'W, 24 
Sep 1999, R..O’Kennon &R. Lance 14463 (UWO);N of Trussville on road to Clay, 1.9 km. after bridge over Highway 
I-59.RHS before 45 mph sign, 33° 40'N, 86° 36'W, 25 Apr 1983, PF. Ulf-Hansen PF029 (UWO). Tuscaloosa Co.: Co. 
Rd. 2,5 km NE of jct. 10 or 51, 33° 12'N, 87° 43'W, 13 Apr 1999, J.B. Phipps 7887 (UWO); probably Co. Rd. 19 (but 
maybe Co.Rd.51),4 km SE of jct.2, 33° 07'N, 87° 44'W, 13 Apr 1999, J.B. Phipps 7883 (UWO); Co. Rd. 2 at 11 km NE 
of jct.51 at overlook, 33° 13'N, 87 ae W, 23 Sep 1999, J.B. Phipps & RJ. oe 8069 (UWO). 

MISSISSIPPI. Noxubee Co.: Butler Rd., running N from MS 21 ca.1 mi W of Gholson, E side of road, 32° 
57'N, 88° 45'W, 28 Sep 1999, J.B. ae RJ. O’Kennon & R. Lance 8103 Ae 


Crataegus pallens Beadle (placed with C. venusta in text) 
ALABAMA. Cherokee Co.: Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18'N, 85° 42'W, 
18 Apr 1999, 1B. Pues & RJ. O’Kennon 7943 (UWO), Dallas Co.: Dallas Co. Rd. 407 between Shephardville and 
Elm Bluff ne ea on right,along NW trending track at 1 km,32° 11'N,87° 05'W, 14 Apr 1999, J.B. Phipps 
7902 (UWO),. ae Co.: banks of Warrior R., 1895, C. Mohr s.n. (ALA). 
NORTH CAROLINA. Buncombe Co.: Biltmore, 18 Sep 1902, Biltmore Herb. s.n. (DOV). 


Crataegus pinetorum Beadle 
ALABAMA. De Kalb Co.: De Soto State Park, 25 Sep 2001,R. Lance 2174 (UWO). Jefferson Co.: near Birmingham, 
20 Apr 1900, CS. Sargent s.n 
GEORGIA. Floyd Co.: Harelee Mtn.,07 Oct 1982, J.B.Phipps 5169 (UWO). Jackson Co.: Tallahassee Shoals 
30 Apr 1930, J.M. Reade E8293 (GA) 
MISSISSIPPI. Smith Co.: SE of Lemon, 30 Sep 1999, J.B. Phipps, R.J.O’Kennon & R. Lance 8092 
Lemon, 12 Apr 2000, J.B. Phipps 8122, 8123 (UWO). 


= 
e) 


— 


UWO); SE of 


Crataegus pulcherrima Ashe 
ALABAMA. Butler Co.: Oaky Streak, approx. 1.5 mi W of Co.59,04 Apr 1985,A.R. Diamond 276 (AUA). Cherokee Co.: 


1004 BRIT.ORG/SIDA 22(2) 


Little River Canyon rimroad ca. 1 km direct NW from Daniel's Gap, 34° 18’N 85° 42'W, 22 Sep 1999, J.B Phipps & RJ. 
O’Kennon 8044 (UWO).Choctaw Co.: 9 mi N of Toxey , just N firetower, Iallahatta formation,03 May 1968, S. McDaniel 
10600 (FSU, IBE). Pike Co.: Sandy ‘Big Pocosin} SE Troy and NE Fairgrounds, 10 Sep 1968,R. Kral 33159 (MO). 

ORIDA. Gadsden Co.: Quincy, in woods, 04 Sep 1895, G.V. Nash 2568 (DOV, MSC); River Jct. 10-11 Aug 
1895, G.V.Nash 2377 (DOV);W of property belonging to River Jct. Hunt Club, beyond end of road on right fork off 
Dolan Road, W, SW of River Jct., 22 May 1982, R.K. Godfrey 79819 (UWO); SW of railroad/Mosquito Creek, WSW of 
River Jct., 22 May 1982,R.K. Godfrey 79815 (UWO). Holmes Co.: Route 185, ca. 1.25 km SW of Co. Rd. 181 just (ca 
150 m) NE of brow on SE Side of road, 30° 52'N, 86° 02'W, 14 Aug 1993,J.B. Phipps 6741 (UWO). Jackson Co.: 9 mi 
NW of Marianna, 21 Oct 1955,R.K. Godfrey & H. Kurz 54277 (DUKE). Leon Co.: Tallahassee, ca. 2 mi S of town, 10 Apr 
1966, R.A. Norris 705 (IBE); woods near Lake Jackson, 07 Apr 1934, L. Griscom 21519 (GH). Liberty Co.: near Bristol, 
06 Sep 1940, WA. Murrill s.n. (GA); near picnic area, Torreya State Park, 16 Jul 1983, R.K. Godfrey 80782 (UWO). 
Wakulla Co.: near Wakulla Springs, 10 Jun 1958, R.K. Godfrey 57030 (DUKE). 

EORGIA. Randolph Co.: Co. Rd. 62,1 3/8 mi from jct. 160, top of bank on LHS, 31° 38'N, 84° 54'W, 03 Oct 
2000, J.B. Phipps & R. Lance 8202 (UWO); Co. Rd. 62, 1 1/8 mi from jct. 160 top of road bank on RHS, 31°38.5'N, 84° 
53.5'W, 03 Oct 2000, J.B. Phipps & R. Lance 8201 (UWO);Co.Rd.62,7/8 mi from jct. 160 top of road bank on RHS, ZA 
38'N, 84° 53'W, 03 Oct 2000, J.B. Phipps & R. Lance 8200 (UWO); SE of Coleman, 6.2 mi from S end of Rd.20 and 1 
mi from jct. with Hwy 160,on E side of road,N 31° 39.468", W 84° 54.642', 27 Sep 2001,R. Lance 2184 (UWO); SE _ 
Coleman, ca.0.5 mi W of Co. Rd. 160,0n N side of Co.Rd.62, across from old drive, N31° 28.630", W 84° 52.875', 27 
Sep 2001,R. Lance 2189 (UWO).Worth Co.: Flint River, 1.9 mi W of Warwick, a 1966, W.R. Faircloth 3901 (GA). 

LOUISIANA. Washington Parish: Bogalusa, 05 Apr 1919, 8.S. Cocks 3149 (NO). 
MISSISSIPPI. George Co.: W bank of Pascagoula R.7 mi E of oe 28 Mar 1975,R.D. Thomas, CM. Allen 

& G. Landry 42999 (NLU). Lauderdale Co.: Mt. Barton and vicinity of S of Meridian, 11 Jun 1957, G.R. Cooley 5274 
(USF). Smith Co.: SE of Lemon on 35.SR 533A, 32° 07'N, 89° 29'W, 26 Sep 1999, J.B. Phipps, R.J.O’Kennon &R. Lance 
8091 (UWO); ESE of Lemon on 533A, right hand side of road at 1.1 km W of jct. 533, 32° 07'N, 89° 26'W, 26 Sep 
1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8095 (UWO); SE of Lemon on 35, SR 355A, 32° 07'N, 89° 28'W, 26 Sep 
1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8093 (UWO), 


~~ 


naa sargentii Beadle 
ALABA Cherokee Co.: Little River Canyon Mouth Park, entrance road, 34° 17'N, 85° 40.5'W, 22 Sep 1999, /.B. 
ie - O’Kennon 8052 (UWO); Little River Canyon, road from canyon mouth to Daniel's Gap, middle se 

17'N, 85° 41'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7932 (UWO); Little River Canyon rimroad ca. 
ee NW from Daniel's Gap, 34° 18'N, 85° 42'W, 22 Sep 1999, J.B. Phipps & RJ. O’Kennon 8041 Tey ced Co.: 
Co. Rd. 15, S of Jackson, 31° 26'N, 87° 52'W, 01 Oct 2000, J.B. Phipps & R. Lance 8194 (UWO). Dale Co.: 3 mi N of 
Clayhatchee, by AL 85,09 Jun 1971, R. Kral 43182 (AUA). Dallas Co.: ca.5 mi E of Sardis on Co. Rd. 30, 32° 15'N, 86° 
48'W, 06 Apr 1984,J.B. Phipps &T.C. Wells 5321 (UWO).De Kalb Co.: De Soto State Park, to rear of campsite #27, 34° 
30.019'N, 85° 37.579'W, 25 Sep 2001, R. Lance 2171 (UWO); De Soto State Park, near campground, 34° 30'N, 85° 
38'W, 08 Oct 1998, J.B. Phipps 7846 (UWO); Lookout Mountain, De Soto State Park, SE side of campground, near 
site # 59,16 Sep 1994, R. Lance 9405 (UWO); Lookout Mountain, De Soto State Park, NW side of campground, near 
site #21, 16 Sep 1994, R. Lance 9402 (UWO); De Soto State Park, campground area, 34° 30”N, 85° 38'W, 22 Sep 
1999, J.B. Phipps & RJ. O’Kennon 8035 (UWO); scrubby woodland on conglomerate ca. 5 km NW of Little R. on 
route 35, 34° 24'N, 85° 39'W,; 08 Oct 1982, J.B. Phipps 5190 (UWO), De Soto State Park, woodland adjacent to 
campground, 34° 30.019'N, 85° 37.579'W, 25 Sep 2001, R. Lance 2169 (UWO); De Soto State Park, campground 
area, 34° 30'N, 85° 38'W, 19 Apr 1999, J.B. Phipps & RJ. O’Kennon 7953 (UWO); Lookout Mountain, back roads S of 
Mentone on route 117, 34° 33"N, 85° 35'W, 08 Oct 1982, J.B. Phipps 5196 (UWO). Marshall Co.: Lake Guntersville 
State Park, Cutchenmire Trail, 34° 22'N, 86° 11'W, 21 Sep 1999, J.B. Phipps & RJ. O’Kennon 8029 (UWO).St. Clair Co.: 
near Springville on U.S.11, 3.5 km from ject. with Old Springville Rd. RHS going N, 50 m. from field edge, along 
fenceline, 33° 46'N, 86° 29'W, 26 Apr 1983, RF UlfHansen PF037 (UWO). Tuscaloosa Co.: Lock 13, 23 May 1952, 
R.L.and O.D.Chermock s.n. (ALA);Kellytown Rd, 1 km from jct.Valley Rd.,33° 07'N,87° 49W, 23 Sep 1999, J.B. Phipps 
& RJ. O’Kennon 8074 (UWO); Co. Rd. 2,5 km NE of jct. 10 or 51, 33° 12'N, 87° 43'W, 13 Apr 1999, J.B. Phipps 7888 
(UWO);Co.Rd. 2 at 11 km NE of jct.51 at overlook, 33° 13'N,87° 42'W, 23 Sep 1999, J.B, Phipps & RJ. O’Kennon 8067 
(UWO); ae Rd. (Co. Rd. 51) at 3.3 km short of Gainesville Rd, 33° 08'N, 86° 44'W, 23 Sep 1999, J.B. Phipps & RJ. 
O’Kennon 8064 (UWOC),. 

FLORIDA. Holmes Co.: Rte. 185, 1.9 km SW of Co. Rd 181, NW side of road, backing onto woodland, 30° 
53'N, 86° 05'W, Apr 1991, 1B. on 6477 (UWO). Leon Co.: Border of second-growth upland woodland, by 
Hartsfield Road, N side, perhaps the equivalent of a city block off capital circle, Tallahassee, 06 Apr 1983, R.K. 
Godfrey 80414 (UWO). 

GEORGIA. Burke Co.: Rte. 56,N of intersection with 80, just N of Lake Crystal Rd.to E, 33° 05'N, 82° 03'W, 09 


— 


— 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 1005 


Apr 1991, J.B. Phipps 6500; (UWO). Floyd Co.: Horseleg Mtn., 34° 14'N, 85° 13'W, 07 Oct 1982, J.B. Phipps 5171 
(UWO). McGee Bend Rd. off Ga. 100 (WSW of Rome), 34° 10.5'N, 85° 22.5'W, 04 Apr 2000, J.B. Phipps 8211 (UWO). 
Horseleg Mtn., near roadside, S-side, 34° 14'N,85° 13'W,07 Oct 1982,/.B. Phipps 5173 (UWO).McGee Bend Rd. off 
GA 100 (WSW of Rome), 34° 10.5'N, 85° 22.5'W, 04 Oct 2000, J.B. Phipps 8210 (UWO). Grady Co.: Upland second 
growth woodland, 3 mi NW of Beachton, by Ga. Rt. 93,31 Mar 1982, R.K. Godfrey 79490 (UWO),. 

MISSISSIPPI. Jasper Co.: 6 mi N of Montrose, 14 Apr 1962, $. McDaniel 2943 (|BE). Noxubee Co.: Butler Rd., 
running N from MS 21,ca. 1 mi W of Gholson, before first side road left, 32° 57"N, 88° 45'W, 16 Apr 1999, J.B. 
Phipps 7914 (UWO). Smith Co.: SE of Lemon on SR 35 to SR 533 at 0.8 km W of bridge, 32° 07'N, 89° 27'W; 26 Sep 
1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8094 (UWO). 


Crataegus tecta Beadle 
ALABAMA. Cherokee Co.: Little River Canyon road, ascent from canyon mouth to Daniel's Gap, ca. halfway up, 
34° 17'N, 85° 41'W, 22 Sep 1999, /.B. Phipps & RJ. O’Kennon 8051 (UWO). Little River Canyon, road from canyon 
mouth to Daniel's Gap, third of way up, 34° 17'N, 85° 41'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7931 (UWO). 
Little River Canyon rimroad,ca.0.6 km N of Daniel's Gap, at overlook, 34° 18'N, 85° 41.5'W, 19 Apr 1999, J.B. Phipps 
& RJ. O’Kennon 7948 (UWO); Little River Canyon rimroad, ca. 1 mi NW of Daniel's Gap, 34° 17'N, 85° 41'W,08 Oct 
1998, J.B. Phipps 7844a (UWO),. Dallas Co.: Elm Bluff, W of Shephardville on Alabama River, wooded slope on N 
side of access road ca. 2 mi back, 32° 11'N,87° 05'W, 02 Oct 1998, J.B. Phipps & R. Lance 7789 (UWO). Geneva Co.: 
Bluff on W side of Hwy 167, near High Bluff and just S of crossing of Chectawhatchee River, 23 Sep 1999,R. Lance 
s.n. (UWO). Houston Co.: Philadelphia, 22 Jun 1984, R.D. Whetstone 14183 (UWO). Montgomery Co.: US 80 at 
Caney Creek, thickets at N side of highway, 32° 19'N,86° 23'W, 24 Sep 1999,RJ.O’Kennon &R. Lance 14451 (UWO),. 
Sumter Co.: 3 mi SW York, 07 Jun 1963, $.B. Jones, Jr. 1319 (GA). Old Bluffport, off S.R.28 ca.7 mi due E of Living- 
ston, along wooded tracks ca. 3/4 mi back from Tombigbee R, 32° 42'N, 88° 07'W, 11 Apr 1998,.8. Phipps 7695 
(UWO). Tuscaloosa Co.: 4.9 mi S of Shirley Bridge (Sipsey) on Co. Rd. 21,07 Apr 1977,J.H. Wiersema 48 (ALU). 4.9 
mi E of Shirley Bridge on Co. Rd. 21,07 Apr 1970, A. Daniels 22 (ALU); Co.Rd. 2 at 11 km NE of jct.51 at overlook, 
33° 13'N 87° 42'W, 23 Sep 1999, J.B. Phipps & RJ. O’Kennon 8066 (UWO); Jct. Kellytown Rd. and Valley Rd., 33° 08'N, 
87° 48'W, st ae 1999, J.B. Phipps & RJ. O’Kennon 8072 (UWO). 

FL A. Gadsden Co.: Turkey Flight Park, Chattahoochee, near parking area, 21 Sep 1999, R. Lance s.n. 


— 


UWO). ite Co.: Three Rivers Recr. Area, 01 Apr 1982, D.S. & H.B. Correll 53541 (USF). Leon Co.: 4 mi E of 
Tallahassee, deciduous woods, 16 Mar 1955,R. K. a 53022 (USF); 2 mi N of Tallahassee, 31 Mar 1958, R.K. 
Godfrey 56323 (NCSC). 
EORGIA. Bartow Co.: US 41 at Pumpkinvine Cr., small flood-plain, 34° 05'N, 87° 44.5'W, 19 Apr 2000, J.B. 
Phipps et al. 8167 (UWO). Clarke Co.: 0.5 mi S of Athens, 26 Apr 1940, WH. Duncan 2073 (GA). Decatur Co.: 
Borders of woodland, between swimming area and Chattahoochee Park, U.S.Corps of Engineers property, close 
to Lake Seminole, ca. 3 mi N of Chattahoochee, Fla., 28 May 1983, R.K. Godfrey and A. Ghalson 80660 (UWO). Early 
Co.: Kolomoki Mounds St. Park woods behind museum, 20 Sep 1999, R. Lance s.n. (UWO); Kolomoki Mounds St. 
Park woods behind museum, 20 Sep 1999,R. Lance s.n. (UWO). Floyd Co.: McGee Bend Rd., off Ga. 100 (WSW of 
Rome), 34° 10.5'N, 85° 22.5'W, 19 Apr 2000, J.B. Phipps et al.8159 (UWO). Randolph Co.: SE of Coleman, ca.0.5 mi 
W of Co.Rd 160,0n N side of Co.Rd.62, across from old drive, N 31° 28.630', W 84° 52.875', 27 Sep 2001, R. Lance 
91 (UWO); SE of Coleman, ca.0.5 miW of Co.Rd 160,on N side of Co.Rd.62, across from old drive, N 31° 28.630', 
W 84° 52.875', 27 Sep 2001, R. Lance 2190 (UWC); SE of Coleman, ca.0.5 mi W of jct. with Hwy 160, 0n S side of 
road, at woods and field corner, near phone box"616-6,"N 31° 39.898', W 84° 53.642', 27 Sep 2001,R. Lance 2186 
(UWO); SE of Coleman, ca. 1.2 miW of Co.Rd 160, on E side of Co. Rd.62,0n bank above road, after creek crossing, 
near eee on hill, os Sep ai R. Lance 2113 (UWO);SE of Coleman,ca.0.5 miW of Ca.Rd 160,0n N side of Co. 
a 630',W 84° 52.875', 09 Apr 2001,R. Lance 2110 (UWO);SE of Coleman, ca.0.1 
mi E of Co. Rd 20, on hilltop seh into young pine plantation, N 31° 38.795',W 84° 55.069", 27 Sep 2001, R. Lance 
2181 (UWO). 
MISSISSIPPI. Lauderdale Co.: Meridian, 15 Apr 1902, T-G. Harbison s.n. (NCU). 
Marion Co.: Pearl River at White Bluff NW of Morgantown, 17 May 1967, S.B. Jones 1236 (NCU).Newton Co.: ca. 2 
mi S of Chunky, Chunky Sand Hill, 32° 18'N, 88° 55'W, 25 Sep 1999,J.B. Phipps, R.J.O’Kennon & R. Lance 8085 (UWO). 
Noxubee Co.: Butler Rd., running N from MS 21 ca. 1 mi W of Gholson, E side of road, 32° 57'N, 88° 45'W, 27 Sep 
1999, J.B. Phipps, RJ. O’Kennon & R. Lance 8097 (UWO). Butler Rd., running N from MS 21 ca. 1 mi W of Gholson, E 
side of road, 32° 57'N, 88° 45'W, 27 Sep 1999, /.B. Phipps, RJ. O’Kennon & R. Lance 8100 (UWO), Butler Rd., running 
N from MS 21 abt 1 mi W of Gholson, just after first side road left, at right, bank, 32° 57'N, 88° 45'W, 16 Apr 1999, 
J.B. Phipps 7917 (UWO),. Pearl River Co.: Off route 43 ca. 12 km S of crossroads (at route 26), 30° 40'N, 89° 45'W, 08 
Apr 1984, 1.B. Phipps &T.C. Wells 5346 (UWO). 
NORTH CAROLINA. Buncombe Co.: Biltmore, 09 May 1902, C.D. Beadle s.n. (DOV). 


1006 BRIT.ORG/SIDA 22(2) 


TENNESSEE. Grundy Co.: NE of Tracy City on mine road from Hobbs Hill to coalmine, 35° 17'N, 85° 34'W, 
01 Sep 1983, PF. Ulf-Hansen PF 112 (UWO) 


Cr Lo venusta Beadle 
ALABAMA. Bibb Co.: Co. Rd. 33, at Cahaba River, 14 Apr 1977,A. Sessler 954 (AUA). Butler Co.: Greenville, - Aug 
en okee Co.: Entrance Road to Little River Canyon Mouth Park, 34° 17"N,88° 40.5'W, 18 
Apr 1999, J.B. Phipps & R.J.O'Kennon 7929 (UWO); Little River Canyon rimroad just NE of Daniel's Gap, 34° 17'N, 85° 
41.5'W,18 Apr 1999,J.B. Phipps & RJ. O’Kennon 7935 (UWO); Little River Canyon rimroad just N of Daniel's Gap, 34° 
17'N, 85° 41.5'W, 22 Sep 1999, J.B. Phipps & RJ. O’Kennon 8049 (UWO); Little River Canyon rimroad just NE of 
Daniel's Gap, 34° 1'N, 85° 41.5'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7934 (UWO); entrance Road to Little 
River Canyon Mouth Park, 34° 17'N, 88° 40.5'W, 18 Apr 1999, J.B. Phipps & RJ. O’Kennon 7925 (UWO), Little River 
Canyon rimroad just NE of Daniel's Gap, 34° 17'N,85° 41.5'W, 18 Apr 1999, .B. Phipps & R.J.O’Kennon 7936 (UWO); 
Little River Canyon rimroad, across canyon of Falls Creek from ‘Pulcherrimae site, NW side of road on fairly straight 
stretch with broad shoulder right, 34° 18.5'N, 85° 41.5'W, 19 Apr 1999, J.B. Phipps & RJ. O’Kennon 7949 (UWO); 
Little River Canyon fea, ascent from canyon mouth to Daniel's Gap, ca. halfway up, 34° 17'N, 85° 41'W; 22 Sep 
1999, J.B. Phipps & RJ.O 8050 (UWO); Little River Canyon Mouth Park, entrance road, 34° 17'N, 88° 40.5'W, 
22 Sep ae Phipps & RJ. O’Kennon 8053 (UWO). Dallas Co.: Dallas Co.Rd.407 between Shephardville and 
Im Bluff near parking area on right, along N-W trending track at 1 km, 32° 11'N, 87° 05'W, 14 Apr 1999, J.B. 
Phipps 7901 (UWO). Marshall Co.: Lake Guntersville State Park on Cutchenmire Trail near Berry Point, 34° 22'N, 
86° 11'W,21 Apr 1999,1.B. Phipps, R.J.O’Kennon & D. Spalding 7966 (UWO); AL 219,4 km S of jct. AL 83 in Talladega 
National Forest, hedgerow on E side opposite two minor roads, 37° 43'N, 87° 06.5'W, 13 Apr 1999, J.B. Phipps 
7882 (UWO). Pickens Co.: dirt road between Fellowship and New Hope churches, 2.2 mi N of Co. Rd. 30,09 Apr 
1977, LJ. Davenport 22 (ALA). St. Clair Co.: ca. 9 mi N Pell City, 22 Apr 1968, R. Kral 30389 (VDB). Tallapoosa Co.: 
Route 50 ca. 1 km E of Martin Dam, 32° 41'N, 85° 53'W, 16 Oct 1984,/.B. Phipps 5507 (UWO). Tuscaloosa Co.: 4.9 
mi S of Shirley Bridge and Co. Rd. 21,07 Apr 1977, 1H. Wiersema 48 (ALA); Co. Rd. 2,10 km NE of jct.10 or 51, SE 
side of, at gate with big overview opposite, 33° 13'N, 87° 42'W, 13 Apr 1999, J.B. Phipps 7891 (UWO). 

FLORIDA. Jefferson Co.: By U.S.Rt.27, 1 mi SE of Jefferson-Leon Co. line and NW of Wacissa-Lloyd Cross- 
road, 27 Mar 1982,R.K. Godfrey 79486 (UWO). 

GEORGIA. Randolph Co.: SE of Coleman, ca.0.5 miW of Co.Rd.160, on N side of Co. Rd.62, across from old 
drive, N 31° 28.630', W 84° 52.875', 27 Sep 2001,R. Lance 2192 (UWO); SE of Coleman, ca.0.5 mi W of Co.Rd. 160, on 
N side of Co. Rd. 62, across i old drive, N 31° Cs 630',W a 52.875',09 Apr 2001,R. ae (UWO), SE of 
Coleman, ca.0.1 mi E of Co.Rd nhillt plantation, N 31° 38.795',W 84° 55.069", 10 Apr 
2001,R. Lance 2115 (UWO);SE cere ca.6.2 mi fon Send of Rd. 20 and 1.6 mi from jct. with Hwy 160,N 31° 
39.468',W 84° 54. On, 10 Apr 2001,R. Lance 2118 (UWO),. 

LOUSIANA.B ille Parish: Bear Creek Gas Field in T16N, R6W, Sec.1 approx.4 mi SE of Bryceland, 12 Apr 
1975, D. Moreland 576 (UWO). 

MISSISSIPPI. Jasper Co.: 6 mi N of Montrose, 20 Aug 1961, §. McDaniel 2683 (IBE). Lauderdale Co.: ca. 2 
mi S of Meridian (T15N R16E S6 NE 4) off side of NW-SE gravel rd., 18 May 1985, TE. Smith et al. 1327 ae 

mper Co,: Few mi along paved road W from MS 39 to Sciple’s Mill turn-off, 32° 52'N, 88° 44'W, 16 Apr 1999, J.B. 
ane 7913 (UWO),. Noxubee Co.: MS 21,5 side, 6.45 km W of jct.39, 32° 58’N 88° 40'W, 16 Apr 1999, 1.B. Phipps 
7918 (UWO). 

TEXAS. Newton Co.: ca. 20 mi NW of Burkville on Little Cow Creek, 11 May 1956, F. Whitehouse 31248 
(SMU). Jasper Co.: 7.5 mi N of Jasper, 11 May 1945, R. McVaugh 6828 (SMU). 


z 


Crataegus ser. Pulcherrimae (Beadle ex EJ. Palmer) Robertson, not assigned to species 
ALABAMA. Cherokee Co.: Little River Canyon Mouth Park, entrance road, 34° 17'N,85° 40.5'W, 22 Sep 1999, /.B. 
Phipps & RJ.O'Kennon 8054 (UWO). Dallas Co.: 9 km E of Sardis on Co. Rd. 30, 32° 15'N, 86° 48'W, 15 Oct 1984, /.B. 
ieee 5485 (UWO). De Kalb Co.: Little River Canyon rimroad, ca. 1 mi NW of Daniel's Gap, 34° V7'N, 85° 41'W, 08 
1998, J.B. Phipps 78446 (UWO). Sumter Co.: Old rane lu S.R. 28 ca. 7 mi due E of Livingston, along 

eee tracks ca. 3/4 mi back from Tombigbee R., 32° 42'N, 88° 07'W, 11 Apr 1998, J.B. Phipps 7697 (UWO). 
Tuscaloosa Co.: Tuscaloosa, 15 May 1874, R.A. Smith s.n. (ALU): aie Co. Rd. 2, hilltop S of Romulus, wooded 
Slope on E side, 10 Apr 1999, R. Lance s.n. (UWO). 

FLORIDA. Gadsden Co.: ae Flight Park, Chattahoochee, 21 Sep 1999, R. Lance s.n. (UWO). Leon Co.: 
woods near L. Jackson, 04 Apr 1934, L. Griscom 21519 (GH). 

GEORGIA. Early Co.: nae Mounds State Park, ca.0.25 mi NE of Tall Pines campground on ridge in 
mixed hardwood forest, 27 Sep 2001, R. Lance 2178 (UWO). Randolph Co.: Co. Rd. 160, just S of Coleman, 31° 


PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 1007 


39.5'N, 84° 53'W, 03 Oct 2000, J.B. Phipps & R. Lance 8197 (UWO); SE of Coleman, ca. 0.5 mi W of Co. Rd. 160,0n N 
side of Co. Rd. 62, across from old drive, N 31° 28.630", W 84° 52.875', 27 Sep 2001, R. Lance 2193 (UWO). 
LOUISIANA. Washington Parish: woods beside Silver Creek and La.438 N of Mt. Hermon, Sec.41, 11S, R9E, 
31 Oct 1980, R.D. Thomas et al 74575 (UWC). 
MISSISSIPPI. Greene Co.: roadside in wooded country near McLain on US 98, 31° 07'N, 88° 48'W, 07 Apr 
1984, /.B. Phipps & T.C. Wells 5334 (UWO). 


ACKNOWLEDGMENTS 


The curators of the following herbaria are thanked for the loan of specimens of series 
Pulcherrimae: A, ALU, AUA, BM, DOV, DUKE, FLAS, FSU, GA, GH, IBE, JSU, LAF LSU, MO, 
MSC, NATC, NCSC, NLU, NO, NY, UNA, US, USCH, USF VDB. Ron Lance, of Chimney Rock 
Park, North Carolina, is thanked for his participation in some of the first author's field trips 
to the area and for his suggestions of sites worth visiting. Susan Laurie-Bourque and Ryan 
Bremmer are thanked for the line illustrations. This work has been funded by a National 
Sciences and Engineering Research Council of Canada Discovery Grant to the first author. 


REFERENCES 


Beapte, C.D. 1902. New species of thorn from the southeastern states, Il. Biltmore Bot. Stud. 1:51-137. 

Kurz, H.and R.K. Goorrey. 1962. Trees of northern Florida. U. Florida Press, Gainesville. 

Lonatey, A.E.1924. Cytological studies in the genus Crataegus. Amer. J. Bot. 11:295-317. 

Pamer, E.J. 1925. Synopsis of North American Crataegus. J.Arnold Arbor. 6:5-128. 

Puipes, J.B. 1988. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. Introduction 
and series Aestivales. J. Arnold Arbor.69:401-431. 

PHipps, J.B. 2005. A review of hybridization in North American hawthorns—another look at “The 
Crataegus problem’ Ann. Missouri Bot. Gard. 92:113-126. 

Pxipes, J.B.and K. Dvorsky. 2006. Crataegus series Parvifoliae and its putative hybrids in the southeast- 
ern United States. Sida 22:423-445 

Reaver, R., J.S. Radrord, and H. LietH. 1974. Modeling important phytophenological events in eastern 
North America. In: H. Lieth. Phenology and seasonal modeling. Springer Verlag, Berlin. Pp. 329- 
342. 

RoserTSON, K.R. 1974. Crataegus ser. Pulcherrimae in, Rosaceae of the southeastern United States. J. 
Armold Arbor. 55:628. 

SarGENT, C.S. 1905. Manual of the trees of North America, Il. Houghton Mifflin, Boston. 

Small, J.K. 1903. Flora of the southeastern United States. ed. 2. Published by the author, New York. 

Tacvent, N.and T.D. Dickinson. 2005. Polyploidy in Crataegus and Mespilus: evolutionary inferences from 
flow ctometry of nuclear DNA amounts. Canad. J. Bot. 83:1 268-1304. (Rosaceae, Maloideae) 

TipestRoM, |. 1933. Crataegus. In:J.K. Small. Manual of the southeastern flora. Univ. North Carolina Press, 
Chapel Hill. 


1008 BRIT.ORG/SIDA 22(2) 


Book REVIEWS 
Tammi RutH Hartuna. 2000. Growing 101 Herbs That Heal: Gardening Techniques, Reci- 
pes, and Remedies. (ISBN 1-58017-215-6, pbk.). Storey Books, Schoolhouse Road, Pownal, 
VT, 05261, U.S.A. (Orders: wwwstoreybooks.com). $24.95, 250 pp., color photographs, 
color illustrations, 8 1/4" x 10". 


With advice for amateur gardeners and for those interested ina starting a medicinal plant farm, this is an enjoy- 
a and useful guide to growing and using medicinal plants. As stated in the title, 101 medicinal plants are cov- 
ered, including f. has Saint John’s Wort (Hypericum perforatum), Echinacea (Echinacea sp.), and lico- 
rice (Glycyrrhiza glabra). Lesser known medicinal plants are also included, such as pleurisy root (Asclepias 
tuberosa). The author is an expert in the uses and cultivation of medicinal plants and is a certified medical herb- 
alist and organic grower. The practical advice in this book is colored by her experiences r unning the Desert Can- 

n Farm & Learning Center in Colorado with her husband. The latter part of the book is dedicated to detailed 
eeu of the medicinal plants and is illustrated with color photographs. Earlier pene include useful 
advice on organic gardening with tips on garden pest control, starting seeds, year-round garden maintenance, 
and composting. An excellent section covers plant selection and garden design. In addition, there are chapters 
dedicated to recipes, as well as instructions on making herbal medicines.—Marissa Oppel, MS, Collections 
Research Assistant, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


JOHANNES SEIDEMANN. 2005. World Spice Plants: Economic Usage, Botany, Taxonomy: teen 


3-540-22279-0, hbk.). Springer-Verlag, Berlin, G wwwspring 
$149.00, 592 pp., 93 color photographs, 91/2" x 10 1 Fa 
This encyclopedic guidebook covers over 1,400 spice plants from the world over, including economically impor- 


tant spice plants and aromatic plants. The lay-out of the book is simple and easy-to-use. The spice plant entries 
are in order alphabetically by scientific name. Each entry includes information on botanical po menc: ature, such 
] 


as family names, species, varieties, subspecies, authorities, synonyms 

including Russian, English, Spanish, Chinese, Arabic, and Japanese, to name a few. Information on uses, eh as 

flavoring, essential oil, condiment, and curry ingredient are included, as well as the part(s) of the plant that is 
o 


used. Information on geographical distribution, including the locations where plant is cultivated is also in- 
f | 


cluded. The extensive reference section is one of the more useful features. It i for those who don't 
want to wade through oceans of a sa oe databases. o references include both primary and s 
ondary scientific Precatire Tl lor pt hs of many of the plants grow- 


ing in the field and in the form in which they are used, such as dried ok T if elf bAGIGES include an index 
co common narnesena an anes of illustrations. Overall, this i book for school 
Mart Issd Oppel, MS, Collecti S TR ch Assistant, Bot -alRes ¢c} 


Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


SIDA 22(2): 1008. 2006 


CRATAEGUS SERIES BRACTEATAE AND TRIFLORAE (ROSACEAE) 


J.B. Phipps R. Lance 
Department of Biology Chimney Rock Park 
A University of Western Ontario Highway 64/74A 
ondon, Ontario, N6A 5B7, CANADA Chimney Rock, North Carolina 28720, U.S.A. 
jphipps@uwo.ca 
K.A. Dvorsky 


Dept. of Biology 
he University of Western Ontario 
don, Ontario, N6A SB7, CANADA 


ABSTRACT 


Crataegus series Bracteatae and a rae, both of the southeastern United States, are revised and shown to pos- 
sess significant similarities. In each series, one of the two species recognized is moderately common, the other 
possibly extinct in the wild, raising conservation concerns. a oe to series is provided and full descriptions for 
both series. Each series has a key to species, detai and county level distribution maps for 


each species, these being the we such known to be produced In addinos each species except C. har bisonii (ser. 


Beacicntiey whichh illustration in Lance and Phipps (2000), is erence 1 eth new line illustra- 


tion. All species are typified and representative specimens are cited for each. 
Key Worps: Crataegus, ser. Bracteatae, ser. Iriflorae, taxonomic revision, typification, illustrations, maps, keys, 


n 


pe 


RESUMEN 


Crataegus series Bracteatae y Triflorae, ambas del sudeste de los Estados Unidos, se revisaron y mostraron tener 
smilies ae En cada serie, una de las dos especies reconocidas es relativamente comun, la otra 
posiblemente extinta en la naturaleza y presentando problemas de conservacion. Se ofrece una clave para las 
series y una dedeapeon cOmIpIa.< de auibes aus una ae las series tiene una a clave de Specs desehipatonies 
detalladas de las especies, y z 
quese prose: esto. jbo se oe una ilustracién todas | peci pt e har bisonii (ser. Bractenlae) 
Lance y Phipps (2000). Se tipifi das | i i 


x .s ps 


eer oneative: de cada una de ellas. 


This paper constitutes another contribution to the continuing series by the first author 
aimed at revising much of Crataegus for the southeastern United States. Other numbers 
have dealt with ser. Aestivales (Phipps 1988), series Apiifoliae, Cordatae, Microcarpae and 
Brachyacanthae (Phipps 1998), Parvifoliae (Phipps & Dvorsky 2006) and Pulcherrimae 
(Phipps et al. 2006). 

This paper is primarily a revision of the species in ser. Bracteatae and Triflorae but 
the series are brought together for convenience and because they do share certain diag- 
nostic characteristics. Appropriate molecular analysis will answer questions about se- 
rial relationships. An abbreviated key to series that occur in the southeastern United States 
(see taxonomic part) illustrates their diagnostic characteristics and separates the two se- 
ries from the others in that region. It is possible that the two series should be fused but it 
is not the intention of this paper to develop that issue. Interestingly though, L. Echols, RL 
and JBP have recently encountered a population in Houston Co., Georgia that appears to 
share cl teristics between C. triflora and C. ashei although on the small sample avail- 
able judgement is withheld. 


SIDA 22(2): 1009- 1025. 2006 


1010 BRIT.ORG/SIDA 22(2) 


Series Triflorae and Bracteatae are both marked by a high degree of glandularity, 
considerable pubescence, unlobed to only very shallowly lobed leaves, few-flowered in- 
florescences, calyx lobe margins dissected and large flowers (20-30 mm diameter). 
Crataegus triflora has an unusually high stamen number relative to other members of 
the genus. Inflorescences borne on the new season’s growth arising direct from woody 
extension shoots may occur in all species except C. ashei and are another attribute of 
both series. A separate discussion of this last point follows below. Elsewhere, this feature 
has only been recorded in Mespilus canescens among closely related taxa. All species are 
bushes or occasionally small trees that are found in open woodland and woodland mar- 
gin often on black-soil prairies. Each series has two species, one moderately common, the 
other very rare or extinct. 

Crataegus generally bears its reproductive organs terminally on woody short shoots. 
Although internode length varies somewhat in these sort shoots, it is shorter to much 
shorter than on extension shoots. Moreover, such short shoots are perennial, borne lat- 
eral to extension shoots, are woody and do not themselves branch. 

The approach used in this paper is essentially the same as that used in others of the 
set. Loans were requested from nearly 40 southeastern herbaria plus three of national 
scope: HUH, MO and US, but material of the two series turned up in only 22 loans. The 
loan material was supplemented by extensive collecting over a number of years mainly 
by JBP and RL. Four phenetic species were recognized, each of which proved possible to 
key out to species named in Beadle (1903) and were typified as appropriate. Detailed se- 
ries and species descriptions were drawn up, as were keys to both series and species. Line 
illustrations for three of the species, county distribution maps and citation of represen- 
tative specimens for all species were then prepared. This will be the first time that any of 
these species has been mapped in detail and we believe that this also is the case for two of 
the species illustrated. 

Species concepts are morphological and based on a cluster of diagnostic characters 
for each taxon and unambiguous separation between each. In this respect it is worth no- 
ticing that, unusually for Crataegus, none of the taxa treated here have synonyms. This 
itself gives a measure of support to our species limits. No explicit data on breeding sys- 
tem was available for any species but recently Talent and Dickinson (2005) have pro- 
duced an array of ploidy level data for taxa treated here, mainly derived from Lance col- 
lections. This did not help in arriving at our specific limits but has potential implications 
for a better understanding of the two series and relevant discussion is provided under the 
species treatments. 


TAXONOMY 


This part of the paper commences witha key designed to discriminate series Bracteatae 
and Triflorae from other sympatric series. It is followed by a conventional treatment of 
the two series and their constituent species with some discussion of each. The abbrevia- 
tion LII refers to the leaf incision index or degree to which sinuses extend toward the 
midvein. Thus, an LIl of 0% represents no lobing, 100% cut to the midvein. 


ABBREVIATED KEY TO SERIES OF CRATAEGUS IN THE SOUTHEASTERN UNITED STATES 


1. Short-shoot leaves lobed with veins to larger sinuses ser. Apiifoliae*, Cordatae*, Microcarpae* 
1. Short-shoot leaves lobed or not, never any veins to sinuses. 
2. Fruit black; short-shoot leaves lacking lobes; thorns < 1cm long, recurved ser. Brevispinae* 


PHIPPS ET AL 1011 


2. Fruit red to yellow; short-shoot leaves lobed or not; thorns mostly 2-5 cm long (except some 
ser. Lacrimatae with very short thorns), straight to slightly recurved (sect. Coccineae, sens. 
lat.). 

3. Ultimate twigs normally not zigzag at any node. 

4. Short-short leaves usually < 1.6 X as long as wide; usually with 2-4 clearly defined 
lobes per side 
5. All green plant parts highly glandular; flowers 20-30 mm wide; inflorescences few- 


flowered; stamens 10 or 30-45 ser. Triflorae 
5. Only some green plant parts soe flowers 12-20 mm wide; inflorescences 
mainly 5-15-flowered; stamens ca. 10 or ca. 20 sect. Coccineae, sens. str.(ser. Molles, 


ccineae, Tenuifoliae, Sie Pruinosae, Intricatae, Pulcherrimae*, C. flava) 
4. Short-short ee mostly 1.75-2 X as long as wide; usually quite unlobed, thou 
sometimes with 1 —2 very small notches per side or alter nati ely with somewhat undu- 


late sides. 
6. Green plant parts mainly + eglandular ser. Aestivales*, Crus-galli, Punctatae 
6. Green plant parts highly glandular. 
7. Leaves glossy, = coriaceous at maturit) ery short-petiolate, 1-4. cm long. 
8 Fruit yellow to ee calyx lobes as long as eect flowers small (12-15 mm 
wide); leaves 1-2 cm long ser. Parvifoliae* 
8, Fruit red; calyx a much shorter than petals; flowers larger (15-20 mm. wide); 
leaves 2-4.cm long ser. Bracteatae 
7. Leaves + matte, usually thin or at least not marked iaceous at maturity,long- 
petiolate; generally 3-6 cm wide and 3-8 cm long at aatcney C. triflora 
(ser. Triflorae, p.p.) 
3. Ultimate twigs always evidently zigzag at nodes ser. Lacrimatae, sens. lat. 


* treated in other papers in this series 


DIRECT COMPARISON OF SERIES BRACTEATAE AND TRIFLORAE 


1. Flowers (15-)20-25 mm wide; stamens ca. 20 or 22—25;leaves + coriaceous at maturity; bushes 
with main stem dominance ser.|. Bracteatae 
1. Flowers 25-30 mm wide; stamens 10 or 30-45; leaves generally thin; multi-stemmed bushe 
ser. Il. Triflorae 


series Bracteatae Sarg. ex Rehder, Man. cult. trees, 1940:363. Type species: C. har bisonii Beadle. 
Group without rank Bracteatae Sarg., Silva Suppl. 13:34. 1902. 


Medium-sized shrubs; thorns 2-4 cm long, somewhat stout, dark and + shiny at 2 yrs. 
old. Leaves deciduous, fairly short-petiolate, less than 1/4 length of leaf blade, pubescent, 
glandular-stipitate; blades 2-7 cm long, broad-elliptic to obovate or roughly suborbiculate 
in general shape; usually unlobed, except sometimes with 2-4 lobed apiculi; margins 
regularly and strongly toothed, the teeth glandular and near the base usually stipitate- 
glandular; venation craspedodromous with 5-7 pairs of lateral veins; + appressed-pu- 
bescent above young, persisting or + glabrescent, below hairy mainly on the veins; tex- 
ture coriaceous. Inflorescences 3-12 flowered, sometimes on leafy short shoots of the 
season; branches densely tomentose, bearing semi-persistent, large, ligulate, gland-bor- 
dered, or smaller, caducous, membranous, bracteoles. Flowers 20-25 mm wide; hy- 
panthium externally tomentose; calyx lobes glandular-pectinate; stamens 20-25, anthers 
usually ivory; styles 3-5. Fruits l-few per infructescence, ca. 10-14 mm broad, hairy, red 
to ruddy; calyx lobes long, conspicuous, patento-ref lexed; nutlets 3-5, dorsally furrowed, 
laterally smooth. 

Habitat and Distribution.—This series comprises one fairly common species, 
Crataegus ashei, and one very rare species, C. harbisonii, together occurring from Louisi- 
ana to Alabama in the piedmont and north into Tennessee and the Virginia Blue Ridge. 


1012 BRIT.ORG/SIDA 22(2) 


They are found in brushy places including prairie margins and open woodland. Unequivo- 
cal ploidy levels obtained are all tetraploid. 

Comment.—The Bracteatae are characterized by relatively few-flowered and fairly 
large-flowered inflorescences and possess very glandular petioles, leaf margins, pedicels 
and calyx lobe margins, which parts are also usually very hairy. They share these char- 
acteristics with the remarkable series Triflorae to which they appear to be related. Num- 
ber of flowers per inflorescence is not a good discriminator among the species mentioned. 
Crataegus harbisonii may bear its inflorescences on side-shoots of the season. The leaves 
of ser. Bracteatae have longer petioles than the superficially similar ser. Crus-galli. 


KEY TO SPECIES 


— 


Leaf blades never, or scarcely perceptibly, lobed (LI] O-5%); marginal teeth 1.5 mm long; 2-yr. 
old thorns fine; all inflorescences borne on perennial sour-shoots; bracteoles less persistent, 


metimes atapieno 1.C. ashei 
1. ae larger leaf t idently, if shallowly, lobed (max. LIl ca. 10-15%); marginal teeth 2 mm 
long; 2-yr.old thorns ae stout; inflorescences quite often on leafy side-shoots of the season; 
bracteoles particularly persistent, often subherbaceous 2.C. harbisonii 


Crataceus ole Beadle, Biltmore Bot. Stud. 1:339. 1900. (Figs. 1, 2). Type: U.S.A. ALABAMA. Mont- 
-o.: Montgomery, between the fairgrounds and the river; May 14 1900, C.D. Beadle 2275 (LECTO- 
TYPE ue designated: US 963476; ISOLECTOTYPES: exist but n.v.). 
Bushes to 5 m tall; thorns few to numerous, 2-3 cm long at 2 yr. old, shiny, very dark 
brown to black, somewhat slender, + straight; extending twigs densely pubescent, at 1 yr 
old tan to chestnut, somewhat shiny; older gray. Leaves deciduous; petioles 0.75-1.5 cm 
long, densely pubescent, bearing stipitate glands; blades 3-6 cm long, elliptic; acute to 
subacute at the apex, base cuneate into the winged upper part of the petiole; generally no 
sign of lobing; margins with even, triangular, acute teeth about 1.5 mm long, the teeth 
gland-tipped, those at the base of the blade glands stipitate; venation craspedodromous 
5-7 lateral veins per side; persistently scabrous-hairy above, below thinly pubescent to 
glabrous on the surface, veins hairy; upper surface shiny mid-green. Inflorescences 3-10 
flowered, not on leafy short shoots of the season; branches densely villous, bearing lin- 
ear, herbaceous to membranous, gland-margined bracteoles. Flowers (15-)20-23 mm wide; 
hypanthium externally tomentose; calyx lobes narrow-triangular, 6-7 mm long, abaxially 
pilose, margins glandular-pectinate; petals + circular, white; stamens 20(-25), anthers 
cream(pink); styles 3(-5) with bristly hairs at the base. Fruit 10-14 mm wide, subglobose, 
dense short-pubescent, orange-red to deep red; calyx lobes long, patent-incurved; nut- 
lets 3(-5), dorsally grooved, sides smooth. 

Common name.—Ashe’s hawthorn. 

Habitat and Distribution.—This is a plant of brushy places and open woodland 
understorey occurring from Louisiana to Alabama and is apparently fairly scarce. Re- 
cently (2000) JBP has come across a highly disjunct record for the Virginia Blue Ridge. 

Comment.—Crataegus dshei has quite often been confused with both C. triflora and 
C. harbisonii. It differs substantially from C.triflora in its different growth habit, smaller 
flowers, 20-24 stamens, and unlobed, generally more coriaceous and shiny leaves. It is, 
however, much more similar to C. harbisonii, mainly differing in being smaller in its veg- 
etative parts, lacking reproductive side-shoots of the season and by having quite entire 
leaves—see the key. Crataegus ashei often has a growth habit not unlike generally sym- 
patric C. berberifolia, with + layered branching and smallish leaves. The seven speci- 
mens cited in Talent and Dickinson (2005) are all tetraploid. 


1013 


PHIPPS ET Al 


Fic. 1. Line illustration of C. ashei. Scale bars = 1cm; S. Laurie-Bourque del. 


1014 BRIT.ORG/SIDA 22(2) 


7 KD) eT ? y! “J 
sy we 
rd 
_ € 
ue 
MT ay 
/ ? 
i oF 4 
f 
s a eee 
a 
/ a“ 


= Number of ay i‘ ie 
0 4-8 . 
Crataegus ashei 1-3 = 9-19 {oat 


Fic. 2. County level distribution map of C. ashei. 


os toe harbisonii Beadle, Bot. Gaz. 28:413. 1899. (Fig. 3). Typr: U.S.A. TENNESSEE. Davidson 
».. W Nashville, hilltop in limestone soil, May 291899, T.G. Harbison 181 (LECTOTYPE here designated: US 
a Specimens of [.G. Harbison 18] (= same tree) exist collected at other times. 


Bushes or trees to 8 m tall; extending shoots sparsely pilose; 1 year old shoots becoming 
smooth, chestnut-brown; thorns 3-4 cm long, stout, + straight, glossy dark brown or near 
black. Leaves deciduous; petioles 1/2-1 cm long, thinly pilose, gland-dotted, winged above; 
blades 4-7 cm long, broadly elliptic to ovate in general shape; tip acute, base cuneate; 
entire or with up to 4 very shallow lobes per side; margins strongly (ca. 5 teeth per cm) 
double-serrate with outward pointing teeth, these gland-tipped early; venation 
craspedodromous with 6-7 pairs of lateral veins; rather shiny and coriaceous at matu- 
rity, appressed-pubescent above when young, when mature pubescent on the veins be- 
low. Inflorescences 5-12 flowered, frequently on leafy short shoots of the season; branches 
pilose, bearing 7-18 mm long, 2-4 mm broad, semi-persistent, subherbaceous, glandu- 
lar-pectinate to glandular-serrate bracteoles; anthesis early May in Nashville, Tennessee. 
Flowers 20-25 mm wide; hypanthium pubescent externally; calyx lobes lanceolate, glan- 
dular-laciniate; petals + circular, white; stamens 20, anther color cream to light yellow; 
styles 3-5. Fruit mostly 12-22 mm wide in recently observed living material, + globose, 
pubescent, orange-red to red; calyx lobes semi-persistent, patent-reflexed; nutlets 3-5, 
dorsally furrowed, sides + plane. 
Iconography.—Lance and Phipps (2000). 


PHIPPS ET Al 1015 


Number of Records 
a 


Fic. 3. County level distribution map of C. harbisonii 


Common name.—Harbison’s hawthorn. 

Habitat and Distribution.—This is a very rare, perhaps extinct species from wooded 
hills near Nashville, Tennessee and one locality in Alabama though its most reliable lo- 
cation is in cultivation in North Carolina and elsewhere due to strenuous efforts at propa- 
gation by the second author. 

Comment.—Harbison’s Hawthorn is very similar to C. ashei and could perhaps be 
thought of as a particularly robust form of that species. However, see discussion of the 
former species for differences, which we consider support specific rank. It has also simi- 
larities to C. triflora (ser. Triflorae) but is easily distinguished from that species by char- 
acteristics of growth-habit (see key), by its smaller flowers (except in the Louisiana and 
some Mississippi forms of triflora) with fewer stamens and by the much more coriaceous 
leaves with much larger and more distant teeth and among the most persistent bracteoles 
in the genus, a feature which led to the creation of series Bracteatae. All five specimens 
cited in Talent and Dickinson (2005), all originating from a tiny population in Nashville, 
Tennessee, are tetraploid. 

IL. ser. Triflorae (Beadle ex C.K. Schneid.) Rehder, Man. cult. Trees, 1940:364. Type sprcirs: C. 
triflora Chapm. 
group without rank Triflorae Beadle in J.K. Small, Fl. SE US, 1903:533. 
sect. Triflorae (Beadle) C.K. Schneid., Ill. Handb Laubholzk. 1:797. 1906. 


Medium-sized shrubs; thorns 2-4 cm long, + slender, dark gray or blackish at 2 yrs. old. 


1016 BRIT.ORG/SIDA 22(2) 


Leaves deciduous, petioles distinct, about 1/3 length of leaf blade, pubescent, glandular- 
stipitate; blades 3-8 cm long, elliptic to nearly suborbiculate in general shape; unlobed to 
shallowly 2-4-lobed per side (max. LI] 10-15%); margins regularly and sharply toothed, 
the teeth glandular and near the base usually stipitate-glandular; venation 
craspedodromous with 5-7 pairs of lateral veins; + hairy above until maturity, hairy 
mainly on the veins below; texture usually thin. Inflorescences 2-6(-12) flowered, nor- 
mally on leafy short shoots of the season; branches densely tomentose, intermixed with 
stipitate glands, bearing herbaceous, semi-persistent, large, ligulate, gland-bordered 
bracteoles. Flowers 25-30 mm wide; hypanthium externally tomentose; calyx lobes broad, 
triangular, deeply glandular-serrate, rather foliaceous; stamens ca. 10 or 30-45, anthers 
usually ivory; styles 3-5. Fruits l-few per infructescence, ca. 10-15 mm broad, subglobose 
to obovoid, hairy, red to ruddy; calyx lobes long, conspicuous, patento-reflexed; nutlets 
3-5, dorsally furrowed, laterally smooth. 

Habitat and Distribution —This series comprises one fairly common (Crataegus 
triflora) and one very rare (C. aust romontana) species, occurring from Louisiana to Geor- 
gia in the piedmont and north into Tennessee with an outlier in Arkansas. They are found 
in brushy places including prairie margins and open woodland, as well as being able to 
tolerate considerable shade. 

Comment.—The Triflorae constitute a distinctive series characterized by few-flow- 
ered and large-flowered inflorescences often borne on leafy short shoots of the season as 
wellas very glandular petioles, leaf margins, pedicels and calyx lobe margins, which parts 
are also usually very hairy. The most similar series is ser. Bracteatae. The 30-45 stamens 
of C. triflora are unique in the genus. Also characteristic are the multiple stems lacking 
main trunk dominance and the inflorescences which are quite often being borne on lat- 
eral shoots of the season. 


KEY TO SPECIES 


1. Leaf blades nearly as broad as long, shallowly lobed; stamens 10 1.C.austromontana 
1. Leaf blades generally much longer than broad, seldom lobed; stamens 30-45 2.C. triflora 


Crataegus austromontana Beadle, Bot. Gaz. 28:412. 1899. (Figs. 4, 5).Type: US.A. ALABAMA. De 
Kalb Co. Valley Head, Sep 1899, C.D. Beadle 1288 (LECTOTYPE here designated: DOV). 
Bushes, to 4+ m tall; twigs of the season densely pubescent; 1 year old twigs brown; thorns 
not recorded. Leaves deciduous; petioles about 30% length of blades, densely pubescent, 
glandular; blades 4-7 cm long, broad-ovate to suborbicular in general outline; shallowly 
3(-4)-lobed, largest LIl ca. 15%; tip subacute, the base cuneate tapering into a upper peti- 
olar wing; margins sharply toothed throughout, the teeth gland-tipped; venation 
craspedodromous, ca. 5 veins per side; when young appressed hairy above, especially on 
the veins, below thinly pubescent except on the veins, when older, surfaces generally 
glabrescent. Inflorescences ca. 3-flowered, on leafy short shoots of the season; branches 
dense-tomentose and bearing caducous, linear, membranous, gland-margined bracteoles; 
flowering 2 May 1911 at Valley Head, Alabama. Flowers ca. 25 mm wide; hypanthium 
externally densely tomentose; calyx lobes ca. 7 mm long, narrow triangular, herbaceous, 
margins glandular-pectinate, adaxially pubescent; petals broadly elliptic, white; stamens 
10, anthers cream; styles 3-5. Fruit 12-15 mm wide, + globose, pubescent, red: calyx-lobes 
conspicuous; nutlets 3-5, dorsally grooved, sides plane. 
Common name.—Valley Head hawthorn. 


PHIPPS ET Al 1017 


Fic. 4. Line illustration of C. austromontana. Scale bars = 1cm; S. Laurie-Bourque del. 


1018 BRIT.ORG/SIDA 22(2) 


Number of Records 


[| 0 4-8 
1-3 9-19 


Fic. 5. C ty level distributi p f C. austromontana 


Habitat and Distribution.—This isa very rare or possibly extinct species not collected 
since 1916. Beadle (1899) reported that it occurred throughout the Sand Mountain region 
of Alabama and had also been collected in the Cumberland Mountains and hill country 
of eastern and middle Tennessee. 

Comment.—lIt is generally similar to Crataegus triflora but has broader, generally 
larger and more deeply lobed leaves and only 10 stamens. 


2. Crataegus triflora Chapman, Fl. S US, 2nd Suppl. 1892:684. (Figs. 6, 7). Typr: U.S.A.Gror- 
GIA: ‘Mountains of Georgia,’ flowering specimen, 1883 or earlier, Chapman Herbarium 847 (NEOTYPE here 
designated: A). A quote froma letter on the type sheet suggests the last possible date for this collection. 

Multistemmed shrubs to 4 or 5 m tall; twigs of current season appressed-pubescent, 2 

year old twigs brown; older gray; thorns usually 2-4 cm long, + slender, + straight, dark 

gray or blackish; compound thorns on trunk. Leaves deciduous; petioles about 1/3 length 
of blades, dense pubescent, also stipitate-glandular; blades 3-8(-10) cm long, broadly el- 
liptic in general shape; entire or shallowly and somewhat irregularly 1-3(-4)-lobed per 
side; tip acute; base cuneate, somewhat tapered into the petiole; margins finely serrate 

(ca. 10 teeth per cm), the teeth usually ca. 1 mm long, gland-tipped; venation 

craspedodromous with 5-7 veins per side; appressed-pubescent above and pubescent 

below when young, only veins below pubescent when old; texture thin. Inflorescences 

2-6(-12) flowered branches dense-tomentose, bearing caducous, linear, thin, herbaceous, 

gland-bordered bracteoles; flowering late April-early May. Flowers 25-30 mm wide: hy- 


1019 


PHIPPS ET AL 


ELE a I 


Ay 
EL essere 
PLE < en) 
+} 3 
TA i 


Fic. 6. Line illustration of C. triflora. Scale bars = 1cm; S. Laurie-Bourque del. 


1020 BRIT.ORG/SIDA 22(2) 


Nace, 


a | 
— | ! 


my ee ‘ a 
Ghasiirsy eee 


Number of Records Toh 
|__| 0 il 4-8 Si 


Crataegus triflora 


Fic. 7. County level distribution map of C. triflora 


panthium externally densely tomentose; calyx lobes 8-10 mm long, herbaceous, glandu- 
lar-pectinate, abaxially densely pubescent; petals + circular, white; stamens ca. 30-45 
(47 in Phipps & Herring 8130), anthers cream; styles ca. 4. Fruit ca. 12-15 mm high, 
subglobose to obovoid, pubescent, ruddy to deep or bright red, occasionally coral-red; 
reflexed calyx lobes long, persistent; nutlets dorsally grooved, sides smooth. 

Iconography.—Color illustration: plate 64, Phipps et al. (2003). 

Common name.—Three-flower hawthorn. 

Habitat and Distribution.—This is a local plant known from six southeastern states 
which extends from Arkansas and Louisiana (rare in both) and Mississippi to Georgia, 
having also been found in Tennessee (also rare). It is only rarely locally abundant. It is 
found scattered in gaps in hardwoods, pine forest, cattle-grazed scrub on blackland soils, 
and sometimes prairie margins. Literature records from Virginia are probably C. x vailiae. 

Comment.—The protologue reads ‘cliffs of Coosa R., Rome’, suggesting that this should 
be the type locality but although this became a popular collection site for the species we 
have seen no material from there of appropriate date for lectotypification. Accordingly, 
we neotypily as above. Crataegus triflora is generally an open-grown, multi-stemmed, 
rather lax bush. Its flowers are among the most spectacular of all Crataegus, in size remi- 
niscent of a wild rose. Although so dramatic and distinct a plant, it unaccountably disap- 
peared from floras between Beadle (1903) and Phipps in Wofford (1989). A fairly rare form 
from central Mississippi has rather ovate leaves which may possess a nearly truncate 
base. This quite distinctive morph also possesses much larger marginal leaf teeth than 


PHIPPS ET Al 1021 


the typical form, more like C. ashei, but it has the flowers of C. triflora. Infrequent speci- 
mens with smallish, unlobed leaves may look very like C. collina if vegetative but are 
easily distinguished by their deeply glandular-serrate calyx lobes, and highly glandular 
leaf margins and petioles. Records from Tennessee generally have relatively broader leaves 
with unusually large marginal teeth. Variability in leaf-lobing is also a notable feature 
though so far it has been correlated with nothing else. The discovery of the large stamen 
number of C. triflora is first recorded in Phipps et al. (2003). The inflorescences may be 
much more floriferous then the name suggests. Another interesting feature of C. triflora 
is that it frequently produces inflorescences on fresh 2-5 cm long peduncles of the sea- 
son that arise from woody extension shoots instead of at the tips of woody short shoots 
as is characteristic of most hawthorn species. Such a feature, rare in Crataegus, is univer- 
sal in Mespilus canescens. Ploidy level determinations in Talent and Dickinson (2005) 
suggest that tetraploidy and triploidy are reasonably common but that diploidy is rare in 
C.triflora. The much smaller leaves and flowers of Caldwell parish, Louisiana popula- 
tions as well as the diversity in leaf shape elsewhere, may possibly one day be shown to 
correlate with ploidy level variations but at present this is not detected. 


APPENDIX 


Consolidated database of cited specimens arranged alphabetically by species, state and 
county 


Crataegus ashei Beadle (number of records: 52) 

LABAMA. Autauga Co.: prairie, Jones Bluff, N of Alabama River, S of Peace, 29 May 2000, R. Lance 2096, 209 
2105 (UNC);same loc, May 1996, J. Allison 9114a, 91146, 9116 (GA). Dallas Co.: Old Cahawba Archaeological Park, 
SW of Selma, Old Negro Cemetery, 32°18'N, 87°06'W, 14 Apr 1999, J.B. Phipps 7895 (UWO). Hale Co.: Gallion, 25 
May 1892, C. Mohr s.n. (UNA). Lowndes Co.: NE of White Hall Holy Ground Battlefield Memorial Park, 32°23'N, 
86°44'W, 14 Apr 2000, J.B. Phipps 8139 (UWO); NE of White Hall Holy Ground Battlefield Memorial Park, 32°23'N, 
86°44'W, 14 Apr 2000, J.B. Phipps 8138 (UWO). ca.8 mi W of Gordonville, an N side of Co. Rd. 12, edge of prairie 2 
mi W of curve of Hwy 12 from N to W in section 27, section 28, just E of stream near BM 317.5, 29 May 1999, Rf. 
Lance s.n. (UWO); NW of Hayneville (W of Montgomery) Holy Ground Battlefield Park (U.S. Army Corps. of Engi- 
neers), 28 Apr 1996, J.R. Allison and A. Schotz 9118 (UWO); NW of Hayneville, Holy Ground Battlefield, 2 Apr 1997, 
R. Lance s.n. (UWO); NW of White Hall, Holy Ground Battlefield Memorial Park, 32°23'N, 86°44'W, 11 Apr 1998, J.B. 
Phipps 7692 (UWO); NW of White Hall, Holy Ground Battlefield Memorial Park, 32°23'N, 86°44'W, 10 Apr 1998, J.B. 
Phipps 7676 (UWO); NW of Hayneville (W of Montgomery) Holy Ground Battlefield Park (U.S. Army Corps. of 
Engineers), 1996/09, A. Schotz s.n. (UWO); NW of White Hall, Holy Ground Battlefield Memorial Park, 32°23'N, 
86°44'W, 24 Sep 1996, RJ. O’Kennon and R. Lance 14453 (UWO). Marengo Co.: near Allenville, 14 Apr 1931, EJ. 
Palmer 38697 (MO); Co. Rd. 51, 2.5 km N of Co. Rd. 54, 32°27'N, 87°40'W, 12 Apr 1999, J.B. Phipps 7871 (UWO). 
Montgomery Co.: Montgomery, 23 Sep 1905, 1.G. Harbison 10041 (NCU);N side of US 80 at Caney Cr., to the NE 
of Montgomery airport, ca. 50 ft NW of 8141, visible from W side of cut, 32°19'N, 86°23’, 14 Apr 2000, J.B. Phipps 
8142 (UWO);N side of US 80 at Caney Cr, to the NE of Montgomery airport, ca. 50 ft NW of 8141, visible from W 
side of cut, 32°19'N, 86°23'W, 14 Apr 2000, J.B. Phipps 8143 (UWO);E Montgomery Airport at jet of Caney Cr.and 
US Hwy 80, in utility right-of-way, ca. 200-250 ft E of creek, 7 Apr 1999, R. Lance s.n. (UWO); N side of US80 at 
Caney Creek, N Side of Rd., 32°19'N, 86°23'W, 12 Apr 1999, 1.8. Phipps 7878 (UWO); no locality, 23 Sep 1905, 7.G. 
Harbison 10041 (UNC);N side of US 80 at Caney Creek, 32°18'N,86°23'W, 1 Oct 1998,/.B Phipps and R. Lance 7780 
(UWO); US 80 at Caney Creek, 32°19'N,86°23'W, 24 Sep 1999, RJ. O’Kennon and R. Lance 14452 (UWO); Montgom- 
ery, 14 May 1900, C.D. Beadle 2275 (US); Montgomery, 28 Sep 1899, no collector (US); Montgomery, 8 Aug 1895, 
C.D. Beadle B895 (US); no locality, 14 May 1900, C.D. Beadle 2275 (US). Unknown Co.: Ranger's Station, 23 May 
1925,W.W.Ashes.n. (NCU). 

LOUISIANA. Caldwell Parish: E of Copenhagen and La. 849 and W of Ouachita River S of Columbia, Sec. 
13.T12N, R4E, 29 Jul 1987, R.D. Thomas 101097 (USCH, UWO); E of Copenhagen and La. 849 and W of Ouachita 
River S of Columbia, Sec. 13.T12N, R4E, 7 Aug 1989,R.D. Thomas 111868 (USCH); ca. 2 km due E of Copenhagen, 
92°02'W, 32°02'N, 10 Apr 1984, J.B. Phipps and T.C. Wells 5366 (UWO); ca. 2 km due E of Copenhagen, 92°02'W, 
32°02'N, 10 Apr 1984, J.B. Phipps and T.C. Wells 5369 (UWO); ca. 2 km due E of Copenhagen, 92°02'W, 32°02'N, 10 


1022 BRIT.ORG/SIDA 22(2) 


Apr 1984, J.B. Phipps and T.C. Wells 5367 (UWO);ca.2 km due E of Copenhagen, 92°02'W, 32°02'N, 10 Apr 1984, J.B. 
Phipps and i Wells 5365 (UWO); ca. 2 km due E of Copenhagen, 92°02'W, 32°02'N, 30 Oct 1985, J.B. Phipps and 
PG. Smith 5875 (UWO); E of Popennage and La. 849 and he of Ouachita River S of Columbia, Sec. 13,112N, R4E, 
17 Nov 1985, R.D.Thomas and B. Liles (MD) 95070 (UWO); E of Copenhagen and La. 849 and W of Ouachita River 
S of Columbia, Sec. 13, T12N, RAE, 3 Ae 1982, R.D. Thomas et al. 82032 (UWO). Sabine Parish: Pleasant Hill Rd./ 
Richmond Rad., 27 Mar 1918, A.8. Cocks 3342 (NO). 

MISSISSIPPI. Chickasaw Co.: MS 41 to SE of Natchez Trace Parkway, 34°05'N, 88°52'W, 13 Apr 1998, /.B. 
Phipps 7713 (UWO). Clay Co.: Co. Rd. 83 at trash dump, Kilgore Hills area NE of Sparta, 33°48'N, 88°56'W, 30 Sep 
2000, J.B. Phipps and R. Lance 8189 (UWO); Co. Rd. 83, 0.5 mi beyond trash dump, Kilgore Hills area NE of Sparta, 
33°48'N, 88°56'W, 30 Sep 2000, 1.8. Phipps and R. Lance 8191 (UWO); Co. Rd. 83 mi after trash dump, Kilgore Hills 
area NE of Sparta, 33°48'N, 88°56'W, 30 Sep 2000, /.B. Phipps and R. Lance 8192 (UWO); Kilgore Hills, on SE side of 
gravel Rd. 83, NE of Sparta (eastbound gravel rd. off Hwy 389 just N of Sparta), just NE of Chickasaw Co. line, S7 
NW4, SW4 1155, R3E, 88°56'W, 1 Jun 2000, R. Lance 00-210 (UWO):; along Co. Rd.8 E Side of Rd, Ca. 3.45 mi E of Co. 
Rd. 389, 33°47'20"N, 38°55'45"W, 01 Sep 2001,R. Lance 2146 (UWO); ca. 4.0 mi SSE of McCondy, just W of NE-SW 
gravel rd.ca.0.5 mi SW of intersection with Hwy 47,115s R4E,$15 SE4, 10 Jul 1985, LE. Smith et al. 1417 (UWO). 
Jasper Co.: ca. 5 mi S of Lake near Tanglewood, N side of Rd., 32°10'N, 89°17'W, 12 Apr 1998, J.B. Phipps 7707 
(UWO). Newton Co.: Black Prairie Region, 4 mi S of Lawrence, 12 Jun 1957, G.R. Cooley and J.D Ray, Junior 5358 
(USF); prairie to S of Moffett Rd. 32°14'N, 89°17'W, 16 Apr 1999, /.B. Phipps 7908 (UWO); E A ee and ca. 
3 mi S of Ponderosa Landing Strip, 32°17'N,89° 1 neo 1998, J.B. Phipps and R.Lance 7811 (UWO); small prairie 
on S side of Moffett Rd. (branching off Gordy Rd.), 32°15'N, 89°16'W, 25 Sep 1999, J.B. Ais pe 8076 (UWO). 
Monroe Co.: Becker Bottom, date and collector missing (IBE). Oktibbeha Co.: Starkville, 4 Sep 1894, Mercer 2813 
(CM). near Clayton Village, just E of MS State (2nd Rd. after Wilkins Rd.), 33°28'N, 88°45'W, 15 Apr 1999, /.B. Phipps 
7905 (UWO),; Clayton Village, E of MS State (2nd Rd after Wilkins Rd.), 33°28'N,88°45'W, 5 Oct 1998, J.B. Phipps and 
R.Lance 7819 (UWO). Scott Co.: N of Sherman Hill, Durand Oak Prairie, 31°17.5'N,89°23'W, 13 Apr 2000, /.B. Phipps 
and J. Herring 8137 (UWO); NE of Sherman Hill, just S of Durand Oak Prairie Forest Rd. 55IF ca.3.5 mi in, 31°17.5'N 
89°23'W, 13 Apr 2000, J.B. Phipps and J. Herring 8134 (UWQ); ca. 5 mi S of Lake near Tanglewood, N side of Rd., 
32°17'N,89° 18'W, 12 Apr 1998, J.B. Phipps 7702 (UWO); NW side of Sherman on FR 551F (gated) off 551, 32°17'N, 

89°22'W, 25 Sep 1999, J.B. Phipps et al. 8081 (UWO).Smith Co.: S of Trenton, Bienville National Forest, 3 Nov. 2005, 
R. — 3025 (NCU). 
IRGINIA. Amherst Co.: Blue Ridge Parkway, along Otter Creek 3.3 mi from Rt. 130, 17 May 1957,R.S. Freer 
24/74 He 


=, 


er austromontana Beadle (number of records: 9) 
MA. De Kalb Co.: Valley Head, 4 Oct 1916, 1G. Harbison 12 (A, NCU); Valley Head, 2 May 1911, 7.G. 
nen 537 (A, CM, UWO); Valley Head, Sep 1899, C.D. Beadle B1288 (DOV); Valley Head, 1911, 7.G. Harbison 3382 
(A); Valley Head, 10 May 1900, C.D. Beadle s.n. (A); Valley Head, 1911, 7G. Harbison 663 (A). 
TENNESSEE. Franklin Co.: near Cowan, 12 May 1900, no collector 42244 (A). 


Crataegus ekg Beadle (number of records: 9) 

ALABAMA e Co.: Gallion, 4 Oct 1898, C. Mohr s.n. (ALU). Montgomery Co.: N side of US 80 at Caney 
Creek, 10 Apr ie Phipps 7674 (UWO); same loc., 1 Oct 1998, R. Lance, s.n.(UWO); same loc., 12 Apr 1999, J.B. 
ice 7879 (UWO). 

TENNESSEE. Davidson Co.: W Nashville, 24-25 Sep 1909, WW. Eggleston 5153 (CM). Obion Co.: near 
Samburg, 26 Jun 1948,A.J. Sharp, E. Clebsch,A. Clebsch, and S. Fairchild 8007 (TENN). Shelby Co.: N edge of Shelby 
Forest, 19 Aug 1947,A.J. Sharp, E.Clebsch,and A. Clebsch 6463 (TENN); roadside, NW of Millington, 29 Jun 1948, AJ. 
Sharp, S. Fairchild and E. Clebsch 8143 (TENN). Weakley Co.: N of Gardner, 23 Jun 1948, S. Fairchild E.Clebsch and A. 
Clebsch 7788 (TENN) 


Crataegus triflora Chapm. (number of records: 133) 

ALABAMA. Autauga Co.: between Jones Bluff and Ivy Creek, S of Peace and N of Alabama River, 29 May 2000, 
R. Lance 2095, 2098, 2100, 2101, 2102, 2103, 2104 (UNC); Jones Bluff, 28 Apr 1996, J. Allison 9114a, 9114b, 9116 (GA). 
Blount Co.: by |-65 ca.5 mi N of Warrior, 23 May 1979,R. Kral63701 (UWO).Calhoun Co.: near Anniston, Area 15D 
Quarry, Ft. McClellan, 22 Apr 1988, R. Smith and D. Spalding 3 (UWO); Fort McClellan Military Reservation, N of 
Anniston, 2 Jul 1998,R. Smith and D. Spalding s.n.(UWO); Fort McClellan Military Reservation N of Anniston 85°46'W, 
33°42'N, 2 Jul 1998, R. Smith and D. Spalding s.n. (UWO); near Anniston Area /SD Quarry [1 5] Fort McClellan 
85°46'W, 33°42'N, 22 Apr 1998,R. Smith and D. Spalding § (UWO); Fort McClellan Military Res N of Anniston 
85°46'W, 33°42'N, 8 Apr 1998, J.B. Phipps 7644 (UWOQ); ca. 4.8 air mi NE of Calhoun Co. anne Anniston, Fort 
McClellan Military Reservation, 13 Aug 1995, JR. Allison et al.8917 (UWO); Fort McClellan Military Reservation 


PHIPPS ET Al 1023 


(old quarry) 33°42'N, 85°46'W, 30 Sep 1998, J.B. bac and R. aes ads (UWO). Cherokee Co.: 1 mi N of Pisgah 
Baptist Church Rd.on Co. Rd. 19,7 May 1974, TA. Atki an n. (ISU). Clarke Co.: dry calcareous 
woods on summit of Salt Mountain, 15 May 1935, RM. Hamper 3374 (A, Ree 15 Apr 1852, Dr. Henry s.n. 
(ALU); Co. Rd. 15, S of Jackson, 31°26'N, 87°52'W, 1 Oct 2000, 1B. Phipps and R. Lance 8193 (UWO). Colbert Co.: 
rocky hillside near Spring Valley, 20 Jul 1899, H. Eggert s.n. (A, MO). Dallas Co.: Pleasant Hill Rd. just before Rich- 
mond Rd., 27 Mar 1918, R.S. Cocks 3342 (NO); Dallas Co. Rd. 407 between Shephardville and Elm Bluff near park- 
ing area on right, 32°11"N,87°05'W, 14 Apr 1999, /.B. Phipps 7898 (UWO);Elm Bluff area ca. 1.5 mi W of Shephardville 
on AL 41 32°11'N, 87°05'W, 14 Apr 1999, /.B. Phipps 7880 (UWO); Elm Bluff area ca. 1.5 mi W of Shephardville on 
AL 41 32°11'N, 87°05! W, 14 Apr 1999, J.B. Phipps 7881 (UWO). Dallas Co.: Rd.407 between Shephardville and Elm 
Bluff near parking on right 32°11'N,87°05'W, 14 Apr 1999,/.B. Phipps 7900 (UWO); Elm Bluff, W of Shephardville 
on abst River, 32°11'N, 87°05'W, 2 Oct 1998, /.B. Phipps and R. Lance, 7790 (UWO); Elm Bluff, near Alabama 
River, 32°11'N, 87°06'W, 2 Oct 1998, 1B. Phipps and R. Lance 7794 (UWO); Elm Bluff, W of Shephardville on Ala- 
bama River, 32°11'N, 87°05'W, 2 Oct 1998, /.B Phipps and R. Lance 7788 (UWO). De Kalb Co.: Valley Head, 4 Oct 

916, 1.G.Harbison 12817 (NCU);Valley Head, Alabama, 2 May 1911,7.G. Harbison 3062 (A);Valley Head, leery slope 
of Lookout Mt., 25 Apr 1924, T.G. Harbison 5906 (A, GH); Collinsville, 26 Sep 1899, no collector (A); sandstone out 
crop on rim of Little aa eee ca.1 miS of Co. Rd.35 on Little River Canyon Rd.,15 Apr 1967,R.C. Clark and K. 
Landers 11174 (UWO). Etowah Co.: on Colvin Mountain (reached by 431 to Cheeseburgh Rd.) 85°52.5'W, 33°68'N, 
8 Apr 1998, .B. Phipps ea (UWO). Franklin Co.: State Rd. 247 at Cedar Creek Public Access Area (Cedar Creek 
Reservoir) 34°33'N,87°58'W,6 Oct 1988,/.B. Phipps and R.Lance 7822 (UWO);AL 247 at Cedar Creek Public Access 
Area, 17 Apr 1999, J.B. Phipps 7919 (UWO); W of Russellville, ca. 1/2 mi S of Co. Rd. 84, ca.0.2 mi N of parking area 
for Cedar Creek Access Area, 200 ft upslope of E bank of Little Rock Creek, 10 Sep 1996,A. Schotz s.n. (UWO); State 
Rd. 247 at Cedar Creek Public Access Area (Cedar Creek Reservoir) 34°33'N, 87°58'W, 7 Oct 1998, J.B. Phipps and 
R.Lance 7823 (UWO). Greene Co.: rich low woods, 1 mi E of Boligee,9 May 1969, .L. Thomas 2058 (ALU). Hale Co.: 
Gallion, Alabama, 17 Apr 1915, 1G. Harbison s.n. (A); E side of AL 69, oak-hickory flatwoods ca. 3/8 mi S of Gallion, 
a little N of Marengo Co. line 32°29'N, 87°42'W, 12 April 1999, /.B. Phipps 7867 (UWO); Gallion, 16 Apr 1915, 1G. 
Harbison s.n. (UNC);E side of AL 69 Oak-hickory flatwoods abt 3/8 mi S of Gallion, a little N of Marengo Co. line, 
32°29'N, 87°42'W, 12 Apr 1999, J.B Phipps 7866 (UWO); 0.3 mi S of Railroad puede hwy 69 just S of Gallion 
Woods on E side of the road, 3 Oct 1988, R. Lance s.n. (UWO). Jefferson Co.: Gate City, near Birmingham, 28 Sep 
1899, C.D. Beadle B1397 (A, ALU, DOV, MO); near Gate City, 5 May 1900, C.D. ee B2280 (ALU, MO);Red Mountain 
Birmingham, 2 Oct 1916, T-G. Harbison 4007 (A); Birmingham, 12 Sep 1911, .G. Harbison 674 (A); Birmingham, 10 
Sep 1911, -G. Harbison 10674 (NCU). Marion Co.: Bear Creek, N of Hackleburg, 28 May 1972, R. Kral 46812 (VDB). 
Marengo Co.: Co. Rd. 51, 2.5 km N of Co. Rd. 54 32°27'N,87°40'W, 12 Apr 1999, .B. Phipps 7869 (UWO); Co. Rd.51, 
2.5 km N of Co. Rd. 54 32°27'N, 87°40'W, 12 Apr 1999, J.B. Phipps 7868 (UWO); Allenville Rd., ca. 1.5 mi N of jct of 
hwy 54, on E side, 9 Apr 1999, R. Lance s.n. (UWO). Montgomery Co.: Black Belt, 19 Apr 1969, 1.C. Chitwood s.n. 
(AUA); Caney Creek, SW of Montgomery, scrubby woodland N side of ae 80 nee W, 32°19'N, 6 Apr 1984, 1B. 
Phipps and T.C. Wells 5312 (UWO);Caney Creek, SW of M ibk 1N side of Hwy 80,86°23'W, 
32°19'N, 6 Apr 1984, J.B. Phipps and T.C. Wells 5313 (UWO); Caney Get SW of Montgomery on US 80, 86°23'W, 
32°19'N, 15 Oct 1984, 1B. Phipps 5482 (UWO); US 80 at Caney Creek 32°19'N, 86°25'W, 24 Sep 1999, RJ. O'Kennon 
and R. Lance 14450 (UWO);N side of US 80 at Caney Creek 32°19’N, 86°23'W, 12 Apr 1999, J.B. Phipps 7877 (UWO); 
N side of US 80 at Caney Creek 32°19'N, 86°23'W, 12 Apr 1999,J.B. Phipps 7876 (UWO);N side of US 80 at Caney 
Creek 32°19'N, 86°23'W, 12 Apr 1999, /.B. Phipps 7875 (UWO);N side of US 80 at Caney Creek to the NE of Mont- 
gomery Airport 32°19'N, 86°23'W, 14 Apr 2000, J.B. Phipps 8141 (UWO); Caney Creek, SW of Montgomery,N side 
of Hwy. 80,86°23'W, 32°19'N, 6 Apr 1984, /.B. Phipps and T.C. Wells 5314 (UWO);N side of US 80 at Caney Creek, to 
NE of Airport 32°19'N, 86°23"W, 10 Apr 1998, J.B. Phipps 7673 (UWO);N side of US 80 at Caney Creek, 32°19'N, 
86°23'W, 30 Sep 1998, J.B. Phipps, and R. Lance 7775 (UWO);N side of US 80 at Caney Creek, 32°19'N, 86°23'W, 1 
Oct 1998, /.B. Phipps, and R. Lance 7779 (UWO). Russell Co.: roadside of Ala.37,ca.4 miS aes 18 Jul 1967, 
R.C. Clark 16135 (NCU). Sumter Co.: Fort Tombigbee, near Epes at Jones Bluff, off Hwy 11 (site owned by Univ. of 
W Alabama) NE of car-park 32°42'N, 88°07'W, 11 Apr 1998, J.B. Phipps 7693 (UWQ); Tombighbee, near Epes at 
Jones Bluff, off Hwy 11 (site owned by Univ. of W Alabama) NE of car-park 32°42'N, 88°07'W, 11 Apr 1998, J.B. 
Phipps 7694 (UWO). Winston Co.: Ranger's Station in Winston Co,, 24 May 1925, WW. Ashe s.n. (NCU). Unknown 
Co.: Ranger's station on a hill, 28 May 1925,W.W.Ashes.n. (NCU); near Nitrate plant, 14 May 1934, CJ. Charlesworth 
and I.G. Harbison 716 (TENN) 

ARKANSAS. Hot Springs Co.: De Roche ce NW 1/4, Sec. 32,T 45,R 19 W, 18 Jul 1977, .L. ee 
427 (UARK); 1.5 mi NW of De Roche off Co.Rd.101,W of S.R. 128,24 Apr 2006,G. Yatski een etal. 06- ie alee WO). 
GEORGIA. Bartow Co.: Pumpkinvine aa 9 May 1996, J. Allison 9138 (GA); Pumpkinvine Creek, 28 Aug. 
1996, J Allison 9431 (GA). Dade Co.: Fox Mountain Track up E side from S of Rising Fawn (at large fa right) 


1024 BRIT.ORG/SIDA 22(2) 


34°44'N, 85°32W, 19 Apr 2000, J.B. Phipps et. al.8153a (UWO); Fox Mountain Track up E side from S of Rising Fawn 
(at large run-off right) 34°44'N, 85°32'W, 19 Apr 2000, J.B. Phipps et al. 8153 (UWO). Floyd Co.: cliffs along S side of 
Coosa River, 3mi SE of Rome, 30 Sep 1951, WH. Duncan 13323 (GA); cliffs of the Coosa River, 6 May 1899, C.S. 
Sargent s.n. (A, MO); cliffs of Coosa River near Rome, no date, I.G. Harbison s.n. (NCU); cliffs of Coosa River near 
Rome, 26 Jul 1899, C.D. Beadle B764 (A); 4 mi below Rome, 6 May 1899, no collector name C-O (A); cliffs of Coosa 
River, Rome, 15 Sep 1899, C.D. Beadle 81184 (A); cliffs of Coosa River, Rome, 20 Sep 1910, 1.G. Harbison 144 (A); cliffs 
of Coosa River near Rome, 5 May 1899, WM. Canby 43 (CM); cliffs of Coosa River near Rome, 5 May 1899, WM 
Canby s.n. (DOV); cliffs of Coosa River near Rome, 1899/9, C.D. Beadle B1397 (DOV); Rome, 1882,A.W. Chapman s.n. 
(GH); McGee Bend Rd, off Ga. 100 (WSW of Rome), 19 Apr 2000, J.B. Phipps et al.8160 (UWO); McGee Bend Rd, off 
Ga. 100 (WSW of Rome), 19 Apr 2000, J.B. Phipps et al. 8156 (UWO); above Coosa River, ca. 3 mi, SW Rome, 4 May 
1980, R. Kral 64929 (VDB). Houston Co.: Oaky Woods Wildlife Management Area, Green Violet Prairie, 14 Apr 
meee Phipps and R. Lance 9064 (UWO); Oaky Woods Wildlife Management Area, Green Violet Prairie, 25 Sep 
05,R. Lance 2930, 2931, 2932 (UWO). Walker Co.: poorly marked trail on Pigeon Mountain from SSW of Marsh 
csi to Bluebird on 34°42.5'N,85°21'W, 18 Apr 2000,/.B. Phipps et. al. | 51 (UWO); Pigeon Mt. near summit, 
e of slope at limestone outcroppings, NE of pocket Brance, N 34° 42.187', W 085° 21.998', 25 May 2000, R. 
en 00-019 (UWO). Unknown Co.: mountains of Georgia, no collector ee eee 1895,A.W. Chapman 
5.n. (DOV). 
LOUISIANA. Caldwell Parish: Ouachita River, ca. 1.25 mi E of Copenhagen, Sec 13,92°97'W, 32°01'N, 20 
Apr Poe F.M. Givens 3604 om gorges E of Copenhagen and La. 849 and W of Ouachita River Sec. B, [ May 
1983, R.D. Thomas and B. Hess 83427 (NLU);clay pine hills along Ouachita River, 1 mi E of Copenhagen Sec. 11,12,13 
on moist sate 17 Mar 1973 PMapeande Jones 422 (NLU); gorges NE of Copenhagen, NE corner of Sec. : clay 
soil, mixed hardwoods forest, 20 May 1973, R.D Thomas, P Marx et al. 3439 & 1042 (NLU); along ridges just W of 
Ouachita River and E of Copenhagen and La.849, 20 Apr 1979,R.D. Thomas and Botany 103 Class 6339 (NLU); hills 
near small creek S of dirt road and E of Copenhagen, Sec. 13, 26 Jun 1973, R.D. Thomas, P Marx and Bot 313 Class 
35892 (NLU); gorges along Creek 1.5 mi E of Copenhagen, 8 Jun 1973,R.D.Thomas and P Marx 35294 & 1323 (NLU); 
open clay pasture NE of Copenhagen NE corner of Sec. 14, 2 Jun 1973, P Marx, PJo nd D.Lawson 1193 (NLU); 
gorges E of Copenhagen and La.849 and W of Ouachita River, Section 13,3 ae 1982,R.D. Thomas, T. Dorris et al 
82032 (NLU); beside small creek W of Ouachita River and E of Copenhagen Sec. 13,6 Aug 1975, R.D. Thomas and 
Bot. 313 class 45887 (NLU); beside small creek W of Ouachita River and E of Copenhagen Sec. 13,6 Aug 1975,R.D 
Thomas and Bot. 313 class 45911 (NLU); 1.5 mi E of Copenhagen, 8 Jun 1973, R.D. Thomas and P Marx 35294 (UWQ); 
1.5 mi E of Copenhagen, 8 Jun 1973, Thomas, R.D. and P Marx 1323 (UWO); E of Copenhagen and La. 849 S of 
Columbia and W of Ouachita River, sec. 13,112 N, R4E, 4 Apr 1987, R.D. Thomas 99232 (UWO; E of Copenhagen 
and La.849 S of Columbia and W of Ouachita River, sec. 13,112 N,R4E, 28 Mar 1987,R.D. Thomas and Bot. 103 class 


99203 (UWO). 
MISSISSIPPI. Chickasaw Co.: ca.6.5 mi SSE of eee) pes Hills, the E edge of the county just W of 
Clay Co. on rural rd, T15S, R3E, $12, the NE/4 of SE/4, 12 Jan 7,J.R. Macdonald et al. 10267 (UWO). Clay Co.: 


Kilgore Hills, ca. 5.5mi NNE Montpelier, NW4 S7, 15 Apr 1981 Pree R. Haynes 24955 (FSU); ca.6.5 mi SSE 
of Houston, a Hills, just inside county from Chickasaw Co.on rural rd,T15S, R4E, S7 the NW/4 of the SW/4, 
12 Apr 1997,.R. MacD ld et al. 10266 (UWO);Co.Rd.83 at trash dump, Kilgore Hills area NE of Sparta, 33°48'N, 
88°56' i: i Sep 2000,J.B. Phipps, and R.Lance 8190 (UWO).Lee Co.: thickets, clay and limestone hills near Verona, 
22 May 1931, EJ. Palmer 39021 (MO). Monroe Co.: S of Amory, ESE of jct. US 45, MS 41,S of 278 at Sandhills Rd., 
33°58'N, 88°34'W, 6 Oct 1998, J.B. Phipps and R. Lance 7820 (UWO). Newton Co.: Eureka Church Cemetery at jct. 
of Gordy Rd.and Moffet Rd. 32°15'12"N,89°15'17"W, 2 Sep 2001,R.Lance 2147 (MO); Eureka Church Prairie jct. of 
Gordy Rd.and Moffet Rd. 32°15'N,89°15'W, 25 Sep 1999, J.B. Phipps, RJ. O’Kennon and R. Lance s.n. (UWO); Eureka 
Church Prairie jct. of Gordy Rd. and Moffet Rd., 32°15'N, 89°15'W, 12 Apr 2000, J.B. Phipps 8124 (UWO); Gordy Rd. 
W of MS 505, NW quadrant, N of Moffet Rd., 32°15'N, 89°16'W, 16 Apr 1999, J.B. Phipps 7917 (UWO); Gordy Rd. W 
of MS 505, NW quadrant, N of Moffet Rd., 32°15'N, 89°16'W, 16 Apr 1999, J.B. Phipps 7910 (UWO); Gordy Rd.W of 
MS 505, NW quadrant, N of Moffet Rd., 32°15'N, 89°16'W,16 Apr 1999, J.B. Phipps 7912 (UWO); intersection of 
Gordy Rd. and Moffett Rd. NW quadrant, 1 km W of MS 505., 32°15'N, 89°16'W, 11 Apr 1999, J.B. Phipps 7862 
UWO); dirt rd. parallel to MS 505 and 2/3 mi W of it, opposite Nance Hill, 32°15'N,89°16'W, 4 Oct 1998, 1.8. Phipps 
and R. Lance 7809 (UWO); dirt rd. parallel to MS 505 and 2/3 mi W of it, opposite Nance Hill, 32°15'N, 89° 16'W, 4 
Oct 1998, J.B. Phipps and R. Lance 7808 (UWO); dirt rd. parallel to MS 505 and 2/3 mi W of it, opposite Nance Hil, 
32°15'N, 89° 16'W, 4 Oct 1998, J.B. Phipps and R. Lance 7807 (UWO); dirt rd. parallel to MS 505 and 2/3 mi W of it, 
opposite Nance Hill., 32°15'N, 89°16'W, 4 Oct 1998, /.B. Phipps and R. Lance 7810 (UWO). Scott Co.: NW side of 
Sherman Hill on 551F (gated) 31°17'N,89°22'W, 25 Sep 1999,/.B. Phipps et.al. 8080 (UWO);W of Forest, S of US 80, 
gated Forest Rd. 537, 32°19.5'N, 89°33'W, 12 Apr 2000, J.B. Phipps 8121 (UWO); NE side of Sherman Hill, just S of 


— 


PHIPPS ET AL 1025 


Durand Oak Prairie, Forest Rd. 551F ca. 3.5 mi in 31°1"N, 89°23'W, 13 Apr 2000, J.B. Phipps and J. Herring 8133 
(UWO);E side of Sherman Hill, edge of Forest Rd. 551F, cutover at ca.0.5 mi in 32°17'N, 89°23'W, 13 Apr 2000, /.B. 
Phipps and J. Herring 8131 (UWO); NE side of Sherman Hill, just S of Durand Oak Prairie, Forest Rd. 551F ca. 3.5 mi 
in, 32°17'N,89°23'W, 13 Apr 2000,J.8. Phipps and J. Herring 8132 (UWO);E side of Sherman Hill, edge of Forest Rd. 
551F,31°17'N,89°21'W, 13 Apr 2000,J.B. Phipps and J. Herring 8129 (UWO);E side of Sherman Hill, edge of Forest 
Rd. 551F, 31°17'N, 89°21'W, 13 Apr 2000, J.B. Phipps and J. Herring 8128 (UWO); E side of Sherman Hill, edge of 
Forest Rd.551F,31°17'N,89°21'W, 13 Apr 2000, J.B. Phipps and J. Herring 8127 (UWO);W of Forest, S of US 80, gated 
Forest Rd.537,0-200 m inside barrier, 32°19.5'N, 89°33'W, 12 Apr 2000, J.B. Phipps 8120 (UWO);W of Forest, S of 
US 80, closed dirt Forest Rd. 537, 32°19.5'N, 89°33'W, 26 Sep 1999, J.B. Phipps 8087 (UWO); Sherman Hill ca.6.5 m 
SW of Lake, 32°17'N, 89°24'W, 4 Oct 1998, J.B. Phipps 7812 (UWO).Tishomingo Co.: Tishomingo aoe 
8 May 1915, T.G. Harbison s.n. (A); Tist Mississippi,8 May 1915, .G.Harbison 12014 (NCU).Webster Co.: NE 
of Maben, CR. 188, 33°34'N, 88°04'W, 27 Sep 1999, J.B. Phipps, RJ. O’Kennon, and R. Lance 8105 (UWO); ca. 2.0 mi N 
of Maben (T20N R12E S20 SW4 NW4) T.E. Smith property, 22 Jun 1985, TE. Smith 1396 (UWO); off MS 15 N of 
Maben (Oktibbeha Co.), first dirt rd. going SE, 33°33'N, 89°04'W, 4 Oct 1998, J.B Phipps and R. Lance 7815 (UWO); 
off MS 15 N of Maben (Oktibbeha Co.), first dirt rd. going SE, 33°33'N, 89°04'W, 4 Oct 1998, 1.8 Phipps and R. Lance 
7816 (UWO); ca. 2.0 mi N of Maben (T.E. Smith property), T20N R12E, 520 SW4 NW4, 2 May 1985, TE. Smith et al. 
1260 (UWO),. 

NORTH CAROLINA. Macon Co.: Chapel Hill, Emilyme Gap, Sep 1912, 1G. Harbison s.n. (NCU); Cullasaja 

River, Aug 1912,W.W.Ashes.n. (NCU). Unknown Co.: Western NC, 23 May 1901,W.W.Ashes.n. (NCU). 


— 


ACKNOWLEDGMENTS 


The curators of the following herbaria: A, ALU, AUA, CM, DOV, FSU, GA, GH, IBE, JSU, 
LYN, MO, NCU, NLU, NO, NY, TENN, UARK, US, USCH, USE VDB are thanked for loan- 
ing specimens for this study. Susan Laurie-Bourque is thanked for her line drawings. Vari- 
ous botanists have provided assistance and companionship in the field, notably RJ. 
O’Kennon. We also want to thank Timothy A. Dickinson and an anonymous reviewer for 
suggesting improvements to the manuscript. 


REFERENCES 


Beabte, C.D. 1903. Crataegus. In: J.K. Small, Flora of the southeastern United States, ed. 2. New York. 

Lance, R.and J.B. PHipes. 2000. Crataegus harbisonii rediscovered and amplified. Castanea 85:291-296. 

Puipps, J.B. 1988. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. Introduction 
and series Aestivales. J. Arnold. Arbor. 69:401-431. 

Pxipps, J.B. 1998. Synopsis of Crataegus series Apiifoliae, Cordatae, Microcarpae and Brevispinae (Ro- 
saceae, Maloideae). Ann. Missouri Bot. Gard. 85:475-491. 

Pxipes, J.B. 1989. Crataegus. In: E.B. Wofford, Flora of the Blue Ridge Mountains. Univ. Georgia Press, 
Athens. 

Pxipps, J.B and K. Dvorsky. 2006. Crataegus series Parvifoliae and its putative interserial hybrids in the 
southeastern United States. Sida 22:423-445, 

Pripes, J.B., R.J.O’KeNNoNn, and R. Lance. 2003. Hawthorns and Medlars. Timber Press. Portland, OR. 

Pripps, J.B.,R.J.O’ KeNNon, and K. Dvorsky.2006. Review of Crataegus series Pulcherrimae (Rosaceae). Sida 
22:973-1007. 

SARGENT, C.S. 1902. Silva of North America, Suppl. XII. Cambridge, Mass 

TatenT, N. and T.A. Dickinson. 2005. Polyploidy in Crataegus and Mespilus (Rosaceae): evolutionary in- 
ferences from flow cytometry of nuclear DNA amounts. Canad. J. Bot. 83:1268-1304 


1026 BRIT.ORG/SIDA 22(2) 


Book NOTICE 


SABEEHA MERCHANT, WINSLOW R. Briccs, and Vick! L. CHANDLER (eds). 2006. Annual Review 
of Plant Biology: Volume 57, 2006. (ISBN 0-8243-0657-0, hbk; ISSN 1040-2519). Annual 
Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. 
(Orders: www.AnnualReviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax; 
onlineactivati lrevi rg). $212.00 (USA), 934 pp., 7 3/8" x 91/4". 


Contents of Volume 57 of Annual Review of Plant Biology 


Looking At Life: From Binoculars to the Electron aa 
MicroRNAs and their Regulatory Roles in Plan 
Cc “hlorophyll pars: Duri ing Senescence 


Quantitative Fluorescence Microscopy: From Art to Science 
Control of the Actin Cytoskeleton in Plant Cell Growth 
Responding to Color: The Regulation of Compl Chromatic Adaptation 


Seasonal Control of Tuberization in Potato: Cxuseroed Elements with the Flowering Response 
Laser Microdissection of Plant Tissue: What You See Is What You Ge 

Integrative Plant Biology: Role of Phloem Long-Distance eens Trafficking 

here Interactions with Plants and Other Organisms 


> 


The role of Root Exudates in Rhizosp 
Genetics of Meiotic Prophase | in Plants 
Biology and Biochemistry of Glucosinolates 
Bioinformatics and its Applications in Plant Biology 
Leaf Hydraulics 
Plant Uncoupling Mitochondrial Proteins 
Genetics and Biochemistry of Seed Flavonoids 
“ytokinins: Activity, Biosynthesis, and Translocation 
Global Studies Cell Type-Specific Gene Expression in Plants 
Mechanism of Leaf-Shape Determination 
Mosses as vee Systems for the Study of Metabolism and Development 


Structure and Function of Photosystems | and II 
Glycosyltransferases of Lipophilic Small Molecules 

Protein Degradation Machineries in Plastids 

Molybdenum Cofactor oo and Molybdenum Enzymes 
Peptide Hormones in Plant 

Sugar sensing and ee in Plants: Conserved and Novel Mech 
Vitamin Synthesis in Plants: i and Carotenoids 


lanisms 


c 


Plastid-to-Nucleus R r etrograde Signalin 
Phe Genetics and Biochemistry of Floral Pigments 
Transcriptional Regulatory Networks in Cellular Responses and Tolerance to Dehydration and Cold Stresses 


Pyrimidine and Purine Biosynthesis and Degradation in Plants 
Phytochrome Structure and Signaling Mechanisms 
Microtubule Dynamics and Organization in the Plant Cortical Array 


SIDA 22(2): 1026. 2006 


MORPHOMETRIC ANALYSIS OF AN AMELANCHIER (ROSACEAE: 
MALOIDEAE) COMPLEX ON THE DELMARVA PENINSULA 
(DELAWARE, MARYLAND AND EASTERN VIRGINIA) 
RESOLVES THE TAXONOMIC IDENTITIES OF 
AMELANCHIER OBOVALIS AND A. CANADENSIS 


Christopher T. Frye 


Maryl iat Depaminent of Natural Resources 
fe and Heritage Service 
‘Netaral Heritage Program 
90 e Mills Road 
Wye Mills, Maryland 21679, U.S.A. 


cfrye@dnrstate.md.us 


ABSTRACT 


Amelanchier obovalis and A. canadensis (Rosaceae: Maloideae) are similar taxa with overlapping geographical 
ranges on the Atlantic Coastal Plain of the United States. Conflicting circumscriptions, aces olen 


tion, and agamosperm y have contributed to the taxonomic confusion in these species. In this study I per- 
form a morphometric examination of nese two species onthe C eastal igh oe Deewale eee and the east- 
ern shore of Virginia—coll Peninsula. 1d tof eleven flowering, fruiting 


and vegetative characters cee from marked individuals. A second dataset contains six minimally correlated 
aes that were obtainable from the type specimen of Amelanchier obovalis. | use Principal Components 
Analysis (PCA) to summarize phenetic similarities and dissimilarities between individuals and agglomerative, 
ierarchical Cluster Analysis to aid in visualization of morphological groups. Selection of groups (putative taxa) 
PCA and 


to inform a vtestabie Sroup, model was Peon? aoe tvery using concordance in phenetic methods 
cl four-group model for which I found a sig- 


iff rall multivariate eff 1 (P Pillai’s Trace, F = 7.29, p> ee < 0.0001) using Multivariate Analysis of Vari- 
ance (MANOVA). Two well- oe groups (1 & 4) corresponded to Amelanchier obovalis and Amelanchier 
canadensis. The remainders of the sample are intermediate between Amelanchier obovalis and Amelanchier 
canadensis and may be the product of recurrent hybridization between these taxa. Lack of clearly delimited 
morphological character space and apparent ecological and genetic were in these taxa suggests the 
taxonomic rank of variety under the earliest combination in Amelanchier, e.g, Amelanchier canadensis var. 
canadensis and Amelanchier canadensis var. obovalis as originally ee by Michaux (1803). I present a taxo- 
nomic treatment and historical analysis of the species taxonomy and a dichotomous key to the varieties. 


RESUMEN 


Amelanchier obovalis y A. ca spiaetiala (Rosaceae: Maloiceae) son taxa psunllatcse con areas eee solapadas 

nlall dos Unidos. I polip oidia, hibridacion 
y ; ae psn nape copula: ala confusién taxonémica en estas especies. En este estudio realizo 
pecies en la parte costera as Delaware, Narang y costa este de Vir- 


1] J a] ct Dp, < g Pee |e oa et omen 


ginia 
de flores, frutos y vase latipes pomaces de ane» pecseeie Un segundo onjinte de datos contiene seis 
écimen tipo de ace CG eae is. es 


-] Caer (c | 


el Analisis Componentes Principales (PC 
y un Analisis de Clusters jerarquizado aes ayudar en la visualiz zai ws grupos eis ase La seleccion de 
jelecde 


grupos (taxa putativos) para informar de un m usando 
conecraanci en los métodos renences (PCA y on ter mg) ye tadistica eecenpie inferencial. analisis da un 
e,F=7.29, 
P > F < 0.0001) usando Andlisis de Varianza Multivariante (MANOVA). Dos grupos muy ek (1 & 4) 
correspondieron a Amelanchier obovalis y Amelanchier canadensis. Los restos de la muestra son intermedios 


SIDA 22(2): 1027- 1048. 2006 


1028 BRIT.ORG/SIDA 22(2) 


— 


entre ‘cious rier apovalisy mmelane/aer Eanes nsis y pueden ser is Progucto de hi bridacion recurrente entre 


estos taxa. | 


} 
y genético en estos taxa en el rango de vate ad bajo la combini acion mas antigua en Amel lamchier, e.g., 
Amelanchier canadensis var. canadensis y Amelanchier canadensi: 


a oe aux aves Presento un tratamiento taxonémico y amalisis historico de la taxonomia de ge especies y 


ra las variedades. 


INTRODUCTION 


Amelanchier Medic. (Rosaceae: Maloideae) is comprised of deciduous small trees and 
shrubs of the North Temperate Zone (Phipps et al. 1990). Amelanchier flowers are pen- 
tamerous, radially symmetrical and protogynous (Robinson 1982; Campbell et al. 1991) 
arranged as inflorescences of simple racemes (except in A. bartraminana). Amelanchier 
fruits are globose, berry-like pomes that attract a wide variety of frugivorous dispersers 
(Robinson 1986). The leaves of Amelanchier, and of the subfamily Maloideae in general, 
vary greatly in size, shape and dentation, and mature foliage on short shoots is often very 
different from those on juvenile (e.g., new sucker growth) or long shoots (Robertson et al. 
1992). The total enumeration of taxa varies among authors. For example, in the north- 
eastern United States, Landry (1975) recognized six species; Fernald (1950a) includes 19 
species but Gleason and Cronquist (1991) only nine species. Synonymy within 
Amelanchier is formidable, reflecting uncertainty surrounding taxonomic concepts of 
species and infraspecific taxa (see Fernald 1941; 1946). Reproductive biology of 
Amelanchier is characterized by polyploidy, agamospermy and extensive hybridization 
between most eastern North American species (Wiegand 1912; 1935; Jones 1946; Cruise 
1964; Campbell et al. 1987; Weber and Campbell 1989; Campbell et al. 1991; Dickinson 
and Campbell 1991; Campbell and Wright 1996; Campbell et al. 1997b; Campbell 1999). 
Recent research has revealed several complex taxonomic issues arising from mating sys- 
tem versatility in the genus including the presence of persistent, morphologically dis- 
tinctive hybrids, e.g. Amelanchier x neglecta (Weber and Campbell 1989; Campbell and 
Wright 1996), microspecies derived from intraspecific hybridization followed by agamo- 
spermy,e.g., Amelanchier “rubra” (Dibble et al. 1998) as well as wide morphological varia- 
tion among populations of single species (Cruise 1964, Campbell et al. 1997a). Reproduc- 
tive versatility in Amelanchier, ie., the combination of vegetative spread, interspecific 
hybridization and formation of agamic complexes has contributed to their phenotypic 
diversity and ecological success, particularly in early successional environments 
(Wiegand 1912, 1935; Cruise 1964; Robertson 1974; Campbell et al 1991; Campbell 1999). 

Despite the abundance and ecological importance of Amelanchier in the southeast- 
ern United States this genus remains poorly studied. This may be due, in part, to the lim- 
ited field work that the primary authors of the genus (e.g., K.M. Wiegand, G.N. Jones, ML. 
Fernald) performed in the southern states but probably more persistently by the diffi- 
culty in gathering adequate material for study, and the limited usefulness of herbarium 
material, including some type specimens, (Wiegand 1912; Nielsen 1939: Fernald 1946; Jones 
1946; Cruise 1964). 

In a long-term study (1999-present), mainly of the stoloniferous members of 
Amelanchier, the most problematic species for confident identification in the field and her- 
barium have been Amelanchier canadensis (L.) Medic. and Amelanchier obovalis (Michx.) 
Ashe. Amelanchier canadensis and A. obovalis occur along the east coast of North America 
with overlapping ranges in the southeastern United States. Herbarium sheets labeled as 
these species often have multiple annotations and the opinion expressed of ten seems cor- 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1029 


related with the general habitat types written on the label and notes on plant habit or height 
noted by the collector and not their morphological characteristics (C. Frye, pers. obs.). 

Three species in addition to Amelanchier obovalis and A. canadensis are listed in flo- 
ras (Tatnall 1946, Brown and Brown 1980) specifically covering the geographic region 
known as the Delmarva Peninsula (Fig. 1), which is comprised of eastern Maryland, Dela- 
ware and eastern Virginia (hereafter referred to as the study area). 

Amelanchier arborea (Michx. f.) Fernald is a species of dry, upland forests that is ex- 
cluded in this analysis. Amelanchier intermedia Spach, is restricted to the Piedmont and 
Mountain physiographic provinces (Wiegand 1920) and was not considered here. 
Amelanchier stolonifera Wieg. is cited by Tatnall (1946) from the Piedmont physiographic 
province in Delaware (not included in the study area). Amelanchier canadensis (L.) Medic. 
var. subintegra Fernald (Fernald 1950b) has been observed by the author in the study 
area but no specimen corresponding to this taxon occurred in the sample. 

Shrubs and small trees that key to Amelanchier canadensis and A. obovalis are wide- 
spread and common in the study area but many characters purported to separate the 
species in taxonomic keys appeared highly variable, temporal, or were the subject of dis- 
agreements among authors. Characters such as plant habit (stoloniferous and colony form- 
ing versus clumped and multi-stemmed), plant height, and ovary pubescence that are 
often used in major couplets to distinguish these species I found were difficult to objec- 
tively determine. For example, nearly all individuals in the study vigorously sucker but 
under certain conditions such as mowing and burning form low, shrubby colonies with 
apparent vegetative spread. Long observation (1999-present) of Amelanchier in the study 
area suggested the presence of at least two coherent taxa; however, several morphologies 
could not be identified with existing taxonomic keys (Wiegand 1912; Fernald 1950a; Rad- 
ford et al. 1968; Gleason & Cronquist 1991). Taxonomic complexity in the study area may 
reflect the presence of numerous distinctive microspecies produced by hybridization 
events followed by asexual seed production and/or cryptic hybridization involving a 
number of agamosperms that would be difficult to detect using morphological charac- 
ters present at a single phenological stage. 

This study aims to determine whether Amelanchier occurring on the Delmarva Pen- 
insula exists as a complex of intergrading, minimally separable phenotypes or contains 
two or more well-defined taxa that are significantly different from each other in mor- 
phology, ecology or both. In order to examine morphological complexity and evaluate 
the various treatments of Amelanchier canadensis and A. obovalis | developed two datasets 
of morphometric characters. The first dataset comprised a “full” set of flowering, fruiting 
and vegetative characters taken from marked individuals (Full dataset). The Full dataset 
was used to explore multivariate relationships between individuals, reduce dimension- 
ality and select taxonomically informative characters. The second dataset comprises a 
reduced set of characters that includes only those characters present on the type speci- 
men of Amelanchier obovalis (Type dataset). The Type dataset is analyzed to determine 
the taxonomic identity of Amelanchier obovalis. A similar analysis using the type of 
ible given the few taxonomically informative char- 


Amelanchier canadensis was not p 
acters present. 

Due to the complex classification history of Amelanchier canadensis and A. obovalis 
I defined groups for analysis a posteriori, without taxonomic classification so as to avoid 
confounding a priori taxonomic identity with the partitioning of morphological varia- 
tion. | use a combination of Cluster Analysis and Principal Components Analysis (PCA) 


BRIT.ORG/SIDA 22(2) 


Fall Line 


1030 
- Western Delaware 
Pee ae Line 
Delaware 
N 
\ . 
Be a 
‘gia 
Maryland |7} _ $ 
y we hath 
rd ae 
« 
Prey 6s 
B { Sussex 
as, 
Fee ts Cath if 
ca nt . 
\ a Dorchester Atlantic 
Le en Ocean 
KE, ry tf Wicomico 
Ob rh a 
— ee 
he Le Ff te 
tr NS oa 
Re oor Seenee me Worcester 
ms PB a r 
» " Fy > ¢ a 


sen 
-Accomack ‘. 
Virginia 2 
Northampton 
Eastem 
US. 
~N 
} 
J 
Fin 1G g phi VW re Let Creal TL Nal D 1 fel W Ic n.l TRI 
r \ - RA | A A rs VV JA I Jal LL ia 7 
y g I 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1031 


to delimit groups (putative taxa) and Multivariate Analysis of Variance (MANOVA) to 
test the best-supported group model. Multivariate analysis of morphological data has 
been used to resolve species complexes in Amelanchier (Dibble et al. 1998), classify taxa 
within groups defined a priori (Saarela et al. 2003; Leonard et al. 2005) and to resolve 
difficulties caused by morphological overlap in infraspecific taxa (Whang et al. 2002). 
The taxonomic treatment and arguments developed herein are presented in a his- 
torical context beginning with André Michaux’s Flora Boreali-Americana (1803) and 
relying heavily on the landmark treatment of Amelanchier by Karl Wiegand (1912). 


METHODS 


Study Site—The Delmarva Peninsula comprises the eastern shore of Maryland, all of 
Delaware (New Castle, Kent and Sussex Counties) and the eastern shore of Virginia 
(Accomack and Northhampton Counties) bounded on the west by the Chesapeake Bay 
and the Susquehanna River and on the east by the Atlantic Ocean (Fig. 1). The Delmarva 
Peninsula lies south of the fall-line, a term used to describe the boundary between the 
Piedmont Province and the Atlantic Coastal Plain and falls entirely within the Outer 
Coastal Plain Mixed Forest Province of Bailey (1995). Elevations on the peninsula are 
mostly less than 20 m ASL and are underlain by relatively young Tertiary and Quater- 
nary sediments of uniformly low relief (Schmidt 1993). It is mostly within the Oak-Pine 
forest region, with the northernmost counties transitional to the Oak-Chestnut region 
(Braun 1950). Significant habitat features include extensive fresh and salt tidal marshes, 
blackwater river swamps, barrier islands and Delmarva Bays (seasonal ponds). The cli- 
mate of Delmarva is moderated by the Delaware Bay, Chesapeake Bay and the Atlantic 
Ocean and is characterized as having cool winters and warm, humid summers. 
Character Selection and Sample Size Determination.— used data from two charac- 
ters: (1) length of the flowering inflorescence [hereafter abbreviated as Flinfllen] and (2) 
length of the lowest pedicel on the flowering raceme [hereafter Flpedlen] as a model for 
determining sample size requirements in the field study. These two characters have been 
used by many authors (Ashe 1903; Jones 1946; Fernald 1941, 1950; Robertson 1974; 
Wiegand 1912, 1920) to describe Amelanchier obovalis and A. canadensis and were readily 
obtainable from herbarium sheets. I took measurements of Flinfllen and Flpedlen from 
54 accessions of both from herbarium specimens borrowed from CM, DOV, FLAS, 
GA, MARY, NA, NCU, PH, US, USEH Vv PL (acronyms follow Holmgren et al. 1990) and of 
personal collections from Delaware, Maryland, North Carolina and Virginia. Flpedlen 
had the lowest dispersion of scores (G = 11.88 mm, o = 5.46 mm, N = 119) and the ratio of 
the standard deviation to the mean, expressed as a percentage, (Coefficient of Variation) 
was 46%. Iterative sample size analysis (Sokal and Rohlf 1968, p. 247) converged on a 
sample size > 53 that would be sufficient for an 80% certainty of determining a 25% dif- 
ference between means at the 5% significance level by Analysis of Variance. For the field 
study I chose an additional thirteen characters that I determined might be informative 
in describing variation in the complex: (1) the proportion of the length of the flowering 
inf] mprised by the lowest flowering pedicel [Flped_flinfl]l, (2) ovary pubes- 
cence during flowering [Flovpubl, (3) flowering sepal position [Flsepos], (4) length of the 
fruiting inflorescence [Frinfllen], (5) length of the lowest pedicel on the fruiting raceme 
[Frpedlen], (6) the proportion of the length of the fruiting inflorescence comprised by 
the lowest fruiting pedicel [Frped_frinfll], (7) ovary pubescence on the developing fruit 
[Frovpub], (8) sepal position on the developing fruit [Frsepos], (9) petal length [Petlen], 


1032 BRIT.ORG/SIDA 22(2) 


(10) petal width [Petwid], (11) the ratio of petal length/petal width [Petl_wratio], (12) the 
leaf length/width ratio[Lfl_wratio]and (13) the ratio of the widest point of the leaf rela- 
tive to total leaf length [Lfwidptl. Table | provides a complete list of characters, their de- 
scriptions and measurements. The means of five replicates per character on each indi- 
vidual were calculated and used in subsequent analyses for all quantitative characters. 
Of the total 15 characters, four were multi-state, categorical variables: flowering and fruit- 
ing sepal position [Flsepos, Frsepos] and flowering and fruiting ovary pubescence 
[Flovpub, Frovpub}| these characters are not included in analysis of variance but were 
collected for descriptive purposes and as a preliminary sample for future analysis. 

Sampling.—During the late winter and early spring | marked 75 individuals of 
Amelanchier with permanent tags on a north-south transect from Cecil County, Mary- 
land (39 32' 38" N) to Northhampton County, Virginia (37 08' 45" N). Additionally, I in- 
cluded a sample from within the core range of Amelanchier obovalis (Bladen County, 
North Carolina) matching the description of Amelanchier obovalis by Fernald (1950a) 
and Radford et al. (1968). I did not attempt to stratify the sample among disturbance lev- 
els, although in order to examine any correlations with disturbance and to avoid ecologi- 
cal bias in selecting the sample I marked individuals in both natural habitats (interior of 
swamp forests, hummocks along fresh-tidal rivers, pine flatwoods) and disturbed habi- 
tats (roadside verges, clearcuts, and powerlines). I returned to each marked individual 
during peak flower and mid-fruiting stages. I defined the peak flowering period as the 
stage when all flowers on the raceme had expanded and mature petals could be easily 
removed from the upper flowers. I defined the early fruiting stage as the period when the 
ovary on the developing fruit had swollen but the fruit was still green or just coloring 
and mature leaves were available. 

Additional effort was made to examine variation in elongation of the lower pedicels 
over time, a character noted by several authors (Ashe 1903; Wiegand 1912; Fernald 1941; 
Jones 1946) and one that I thought might be informative for understanding the various 
taxonomic concepts of Amelanchier obovalis and A. canadensis. This additional charac- 
ter isnot used in the multivariate analysis. lexamined the difference between mean length 
of the lowest pedicel at peak flower and at mid-fruit by analysis of variance. Inflores- 
cence and pedicel measurements were taken in the field, all other measurements were 
taken in the lab. All measurements utilized a digital caliper and an ocular micrometer 
scaled to tenths of a millimeter. Vouchers of flowering and fruiting material were col- 
lected from marked individuals at each site; these are deposited at the Maryland Depart- 
ment of Natural Resources Herbarium (Tawes), Annapolis, MD. Sampling began on 18 
April 2005 and continued through 26 May 2005. Flowering, fruiting and leaf measure- 
ments were successfully gathered for 61 individuals. 

Quantitative Morphological Analyses.—My overall objective for the analysis of quan- 
titative morphological characters is to test the null hypothesis of no significantly distin- 
guishable groups of individuals. To test the null hypothesis I looked for discrete clusters 
using Principal Components Analysis (PCA) based on the correlation matrix of eleven 
(Full dataset) and six (Type dataset) quantitative characters standardized by log trans- 
formation. | also used the Full dataset to reduce the dimensions of the dataset by elimi- 
nating highly correlated characters (Pearson's r >0.7). Proportion of variance explained 
by each axis was compared with that expected under a broken-stick model Jackson 1993). 
I plotted the first principal component against the second to graphically portray any 
relationships among individuals. Additionally, I created a second matrix from an 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 


Taste 1.List of the 15 character acronyms, their descriptions and measurement. All measurements taken to the 


nearest tenth of a millimeter. 


Character acronym Description and Measurement 

Flinfllen Length of the flowering inflorescence at peak flower. Measured from the base 
of the distal-most flower on the raceme to point of insertion on the stem 

Flpedlen Length of the lowest pedicel on the flowering raceme at peak flower. Measured 
from the base of the ovary to point of insertion on the raceme. 

Flped_flinfl Proportion of the length the flowering raceme comprised by the lowest pedicel. 

Flovpub vary pubescence on the terminal flower of the flowering raceme scored from 
1 (glabrous) to 4 (densely tomentose) 

Flsepos Sepal position on the terminal flower of the flowering raceme scored along a scale 
from 1 (erect) to 4 (tightly reflexed) in comparison to an imaginary transverse plane 
through the ovary. 

Frinfllen Length of the fruiting inflorescence. Measured from the base of the distal-most 
fruit on the raceme to point of insertion on the stem 

Frpedlen Length of the lowest pedicel on the fruiting raceme at mid-fruit. Measured from the 


Frped_frinfl 
Frovpub 


base of the ovary to point of insertion on the raceme 

Asem of the length of the fruiting raceme comprised by the lowest pedicel. 
vary pubescence on the terminal fruit of the fruiting raceme scored along a scale 

an 1 (glabrous) to 4 (densely tome 


Nn 
cy 


Frsepos Sepal position on the terminal fruit of the fang raceme scored along a scale from 
1 (erect) to 4 (tightly reflexed) in comparison to an imaginary transverse plane 
through the ovar 

Petlen Length of mature, expanded petals measured from distal-most point to insertion 


1033 


on the hypanthiu 

Width of mature smpahued petals measured at the widest point. 

Ratio of petal length/petal width. 

Lfl_wratio Ratio of leaf length/leaf width. 

Lfwidpt Ratio of the widest point along the leaf axis (measured from base)/total leaf length. 


agglomerative, hierarchical Cluster Analysis using Euclidean distance and Ward’s method 
of linkage (McCune & Grace 2002). This second matrix was used as an overlay on the 
PCA scatterplot to assist in visualizing groups. PCA and Clustering were performed on 
Pcord version 4.14 (McCune & Mefford 1999). 

A separate objective was to determine the a (group membership) of the type 
specimen of Amelanchier obovalis in multivari space. 1 performed both PCA 
and Clustering based on the correlation matrix of six quantitative characters derived 
from analysis of the Full dataset and that were also obtainable from the type specimen: 
Petlen, Flinfllen, Flpedlen, Flped_flinfl, Lfl_wratio, and Lfwidpt. The Type dataset was 
organized into a group by character matrix and exported to SAS version 9.1 (SAS Insti- 
tute, Cary, NC, USA) for Multivariate Analysis of Variance (MANOVA, GLM procedure). 
MANOVA tests for an overall multivariate effect of the group model and performs pair- 
wise comparisons between character means (mean vectors) for all groups for each char- 
acter. Approximate univariate normality of morphometric characters within each group 
was assessed by inspection of frequency histograms, accepting normality with skewness 
< 1 (McCune & Grace 2002). MANOVA is robust to multivariate non-normality and 
Levene’s test was used to ascertain homogeneity of variances within each group. MANOVA 
is used to test the null hypothesis of no significant differences between the mean vectors 
of each group. 


1034 BRIT.ORG/SIDA 22(2) 


Assessment of Taxonomic Rank.—| utilize the concepts of genetic and ecological 
exchangeability introduced by Crandall et al. (2000) to assess population distinctive- 
ness and extend that method to assessing taxonomic rank. Species populations are ge- 
netically exchangeable if there is evidence of ample gene flow between them. Ecological 
exchangeability is demonstrated if individuals can be moved between populations and 
can occupy the same ecological niche. Exchangeability is viewed from both recent and 
historical perspectives, e.g., two taxa may currently experience gene flow due to break- 
down of intrinsic or extrinsic barriers via ant} causes but the populations may 
have historically experienced (e.g., presettlement) little gene flow. 


RESULTS 


Full Dataset.—Eigenvalues for the first two axes, representing 56% of the total variation, 
exceeded their broken-stick values and were chosen for interpretation (Table 2). PCA-1 
comprises > 37% of the variance in the dataset. Five morphological characters ordinate 
individuals along PCA-1. These are, in descending order in the loading matrix (Table 3): 
Flpedlen, Frpedlen, Frinfllen, Flinfllen and Petlen. PCA-1 is thus a size axis. PCA-2 com- 
prises > 19% of the variance in the dataset. Five morphological characters ordinate indi- 
viduals along PCA-2. These are, in descending order in the loading matrix (Table 3): 
Flped_flinfl, Frped_frinfl, Lfl_wratio, Flinfllen and Frinfllen. PCA-2 is chiefly a shape 
axis. The characters most highly correlated with both PCA axes are also very highly cor- 
related with each other, in particular, flowering and fruiting stages of the inflorescence 
are redundant (Table 4). Flowering characters were selected for the final matrix. 

Type Dataset.—Six characters were used in the analysis of the Type dataset: Petlen, 
Flinfllen, Flpedlen, Flped_flinfl, Lf{l_ Wratio and Lf widpt. These six quantitative char- 
acters were obtainable from the type specimen of Amelanchier obovalis and 
serendipitously mirrored the set of minimally correlated characters derived from analy- 
sis of the Full dataset with the exception of Flpedlen (r = 0.708), which I maintained 
because of its use as an informative character in most treatments. The dendrogram pro- 
duced by Cluster Analysis (Fig. 2) is scaled by an objective function that measures the 
loss of information at each step; as groups are fused the amount of information decreases 
until all groups are fused and no information remains (McCune & Grace 2002). In the 
iterative process, five and higher order groups were composed of single (outlier) indi- 
viduals and selection of three groups combined groups 2 and 3 into a single membership 
cluster. I chose to overlay a four-group cluster on the PCA as this appeared to be the maxi- 
mum number of well-supported groups (e.g., long branches of the dendrogram) and cor- 
responded to my field observations of the number of putative morphological forms prior 
to sampling and roughly to the different habitat types that were sampled (e.g., tidal and 
nontidal swamps, low Pine woodlands, and sand ridges). The PCA scatterplot with Clus- 
ter overlay (Fig. 3) shows a single distinct cluster at the far left of PCA-1 and central to 
PCA-2. The eigenvalues for the first two principal components, representing >65% of the 
total variation, were selected for interpretation; however, only the second axis exceeded 
its broken-stick value (Table 5). PCA-1 comprises > 39% of the variance in this dataset. 
Three morphological characters ordinate individuals along PCA-L. These are, in descend- 
ing order in the loading matrix (Table 6): Flpedlen, Flinfllen and Petlen. PCA-1 is a size 
axis clearly separating group | (positive affinities) and group 4 (negative affinities) but 
with overlap between groups 2 and 3 at the origin. PCA-2 comprises > 26% of the vari- 
ance in the dataset. Three characters ordinate individuals along PCA-2. These are, in 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1035 


Tas.e 2. Full dataset. Eigenvalues, percent variance extracted and broken-stick comparison of four Principal 
Component axes with eigenvalues > 1. 


Axis Eigenvalue Percent of variance Cumulative Broken-stick 
Percent Eigenvalue 

1 4.106 37.325 37.325 3.020 

2 2.112 19.204 56.529 2.020 

3 1.482 13.470 69.999 1.520 

4 1.199 10.902 80.901 1.187 


Tasie 3. Full dataset. Principal components loading matrix of the correlations between eleven morphological 


characters and | t scores (eigenvectors). The five highest correlations with the eigenvector are in bold 
type. 
Character Eigenvector 

1 2 
Petlen -0.3267 0.0632 
Petwid -0.2567 0.0249 
Petl_wratio -0.0921 0.0471 
Flinfllen -0.3826 -0.3641 
Flpedlen -0,.4620 0.0131 
Flpedl_flinfl -0.2524 0.4800 
LfL_wratio 0.0045 -0.4525 
Lfwidpt 0.0434 0.3073 
Frinfllen -0.3906 -0.3242 
Frpedlen -0.4433 0.0426 
Frpedl_frinfl -0.2107 0.4732 


descending order in the loading matrix (Table 6): Lfl_wratio, Flped_flinfl and Lf widpt. 
PCA-2 isa shape axis that most clearly separates group 3 (positive affinities) from group 
2 (negative affinities). 

The type specimen of Amelanchier obovalis is nested near the center of group 1 out- 
side of the range of variation (along PCA-1) in other groups (see Fig. 3). Descriptive statis- 
tical analysis of the four-group model, calculating the means, standard deviations and 
90% confidence intervals of morphological characters most highly correlated with PCA- 
l and PCA-2 provide some morphological coherence of the groups with strong positive 
or negative affinities along the principal component axes (Table 7). Group 1 (triangles) 
consists of individuals (on PCA-1) having short petals (6.8 + 0.9mm), short flowering 
inflorescences (28.7 + 0.9mm), and short lower pedicels (7.9 + 1.3 mm). On the second 
PCA axis group | consists of individuals with leaves averaging 1.8 times as long as wide 
(1.79 + 0.22) that are widest above the midpoint (0.548 + 0.037) and have more or less 
cylindrical inf] (0.283 + 0.053). Group 4 (stars) consists of individuals (on PCA- 
1) having long petals (8.8 + 1.2mm), long flowering inflorescences (47.8 + 3.9mm) and 
long lower pedicels (15.8 + 1.7mm). On the second PCA axis group 4 consists of individu- 
als with leaves averaging 1.8 times as long as wide (1.80 + 0.18), that are widest near the 
midpoint (0.519 + 0.034), and have spreading inflorescences (0.335 + 0.042). Group 2 


Taste 4. Full dataset. Cross-products matrix of Pearson's Correlation Coefficients (r) for eleven morphological characters. Values of r 0.7 


= 


r ounded) are in bold type. 


Petl_wratio 
Flinfllen 
Flpedlen 
Fiped|_flinfl 
Lfl_wratio 
Lfwidpt 
Frinfllen 
Froedlen 
Froedi|_frinfl 


Petlen Petwid Petl_wratio Flinfllen Flpedlen Fipedl_flinfl  Lfl_wratio Lfwidpt Frinfllen Frpedlen Frpedl_frinfl 
1 0.669 0.396 0.368 0.497 0.327 0.005 0.137 0.385 0.398 0.156 
1 -0.410 0.328 0.356 0.163 -0.049 0.164 0.314 0.326 0.118 
- 1 0.062 0.185 0.202 0.043 -0.057 0.104 0.107 0.057 
: 1 0.778 -0.018 0.229 -0.271 0.839 0.652 -0.021 
- : = 1 0.612 -0.024 -0.117 0.708 0.794 0.372 
= : : 1 0.329 0.146 0.058 0.434 0.616 
= : : - - 1 -0.168 0.192 -0.039 -0.311 
- ; - 1 -0.179 -0.094 0.073 
- - : - - - 1 0.759 -0.042 
: : - - - - 1 0.602 


9E0L 


(2)@2 VaIs/DYO'LINS 


Q¢ 74 


5.5E-04 6.4E-01 


Distance (Objective Function) 
1.3E+00 


Cluster Analysis 


Information Remaining (%) 
50 


Group 1 


Group 3 


Q20n 
See 
za® 


Group 4 


kings-d 3 
jonmt-b f 
forestin 


X31dWO) YFIHDNVTIWY NV 4O SISATVNY DINLIWOHdYOW ‘JAYS 


ZE0L 


1038 BRIT.ORG/SIDA 22(2) 


Principal Components Analysis 


4 4 Group 
& 1 
¢ 3 
x 4 
J ray O02 
e e 
25 ¢ 
Sa 
. 
* * o K 
7 & 
7k ° 
* my 
“ A 
' * & 
od xe Go a A 
O * ¢ = 
Oo *R O a 
aK oO A 
© & ray 
ray 
, Re TYPE~ A&A 
A 
@ & 
A 
2 “7 oO oO 
A 
JJ © 
A 
-4 
-3 -1 1 3 
Raced 
Fic, 3. PCA scatterplot with Cluster overlay. Symbols d bership: triangl group 1, circles = group 2, diamonds = 


group 3, stars = group 4. Type specimen of Amefanchier sbolle indicated. 


(circles) consists of individuals (on PCA-1) having intermediate petal lengths (7.7 + 0.8 
mm), flowering inflorescences (35.1 + 4.6) and lower pedicels (12.6 + 1.7mm). On the 
second PCA axis group 2 consists of individuals with leaves averaging 1.7 times as long as 
wide (1.67 + 0.14) that are widest above the midpoint (0.547 + 0.057) and have widely 
spreading flowering inflorescences (0.363 + 0.043). Group three (diamonds) consists of 
individuals (on PCA-l) having intermediate length petals (7.3 + 0.9mm), long flowering 
inflorescences (39.6 + 3.0mm) with short lower pedicels (10.8 + 1.1mm). On the second 
PCA axis group 3 consists of individuals with leaves averaging 1.9 times as long as wide 
(1.87 + 0.17), that are widest at the midpoint (0.503 + 0.029), and have cylindrical flower- 
ing inflorescences (0.272 + 0.026). 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1039 


Taste 5. Type dataset. Eigenvalues, percent variance extracted and broken-stick comparison of two Principal 
Component axes with eigenvalues > 1. 


Axis Eigenvalue Percent of variance Cumulative Broken-stick 
Percent Eigenvalue 

1 2.345 39.082 39.082 2450 

2 1.575 26.243 65.325 1.450 


Tape 6. Type dataset. Principal Components loading matrix of the correlations between six morphological 
characters and component scores (eigenvectors). The three highest correlations with the eigenvector are in 
bold type. 


Character Eigenvector 

1 2 
Petlen -0.4524 -0.1036 
Flinfllen -0.4994 0.4079 
Flpedlen -0.6326 0.0032 
Flpedl_flinfl -0.3720 -0.5164 
LfL_wratio 0.0144 0.5610 
Lfwidpt 0.0842 -0.4914 


Multivariate Analysis of Variance.—There is a significant overall multivariate effect 
of the four-group model (Table 8) and significant F ratios were found for five of the six 
quantitative characters (Table 9). Pair-wise comparisons between all groups (Table 9) de- 
termined significant differences between all pairwise group mean vectors for Flinfllen 
and five of six comparisons for Flpedlen. Petlen was significant for 4 of 6 group compari- 
sons, Flped_flinfl was significant in 4 of 6 group comparisons, and Lf widpt was signifi- 
cant for 2 of 6 group comparisons. Differences between group mean vectors for LfL__Wratio 
were not significant. Group 4 contains the most distinctive individuals, as expressed by 
all pairwise comparisons of mean vectors for Petlen, Flinfllen, and Flpedlen and two of 
three pairwise comparisons for Flped_flinfl. Group 1 is also distinctive as expressed by 
all group parisons for Flinfllen and Flpedlen, 2 of 3 pairwise comparisons for Petlen, 
2 of 3 pairwise comparisons for Flped_flinfl,and 1 of 3 pairwise comparisons for Lf widpt. 
Group 2 is significant in all pairwise comparisons for Flinfllen, 2 of 3 comparisons of 
Flpedlen, 2 of 3 pairwise comparisons for Petlen, 2 of 3 pairwise comparisons for 
Flped_flinfl, and 1 of 3 pairwise comparisons for Lf widpt. Group 3 is significant in all 
pairwise comparisons for Flinfllen, 2 of 3 comparisons of Flpedlen, but only 1 of 3 pairwise 
comparisons for Petlen, 2 of 3 comparisons for Flped_flinfl and 2 of 3 comparisons for 
Lfwidpt. 

The presence of a large sample of individuals with intermediate phenotypes sug- 
gests ample gene flow (genetic exchangeability) between the groups. Ecological exchange- 
ability is demonstrated by the occurrence of morphologically distinct individuals oc- 
curring within the same habitat type. 

Taxonomic Usefulness of the Elongation of the Lowest Pedicel.—There is significant 
variation among groups in the elongation of the lowest pedicel from peak flowering to 
mid-fruiting as determined by Analysis of Variance. The difference between the mean 


1040 BRIT.ORG/SIDA 22 


= 


2) 


Taste 7. Descriptive statistics of six characters most highly correlated with the principal components (PCA-1 and 
PCA-2) under a four-group model.Columns under Group labels N = number of individuals in g 

+ standard deviations, (ranges), [minimum, maximum], {90% Conndence ees All mencurements ae 
raw (untransformed) scores. 


Character Group 1 Group 2 Group 3 Group 4 
N=24 N= 13 N=14 N=11 
Petlen 6.8 + 0.9 7.7+08 73+09 8.8 + 1.2 
(3.0) (3.5) (3.0) (4.0) 
[5.6; 8.6] [5.9;9.4] [5.8; 8.8] (6.5; 10.5] 
{6.5—7.1} {7.3—8.1} {6.9—7.7} {8.1—9.5} 
Flinfllen 28.7 +09 35.1446 39.6 + 3.0 478 + 3.9 
(17.3) (14.4) (8.8) (10.6) 
[18.7; 36.0] (28.4; 42.8] (35.0; 43.8] (42.2; 52.8] 
{27.1—30.3} {32.8—37.4} {38.2—41.0} {45.7—49,9} 
Flpedlen 79413 1236617 10.84 1.1 15.8+ 1.7 
(5.7) (5.6) (3.4) (5.2) 
[5.6: 11.3] (9.6; 15.2] (9.3; 12.7] [13.2; 18.4] 
{7.4—8.4} {11.8—13.4} {10.3—11.3} {14.9—16.7} 
Flpedl_flinfl 0.283 + 0.053 0. oe + 0.043 Hees + 0.026 0. ee + 0.042 
(0.238) (0. (0.108) (0.123) 
(0.167; 0.405] (0. ae 0.456] [0. ‘ 2; 0.320] (0. 0.396] 
{0.264—0.302} {0.342—0.384} {0.260—0.284} {0.332—0.378} 
LfL_wratio 1.79 + 0.22 1.67 + 0.14 1.88 + 0.17 1.80+0.18 
(0.70) (0.45) (0.58) (0.59) 
(1.47; 2.17] [1.41; 1.86] (1.64; 2.22] [1.56; 2.15] 
{1.71—1.87} {1.60—1.74} {1.80—1.96} {1.70—1.90} 
Lfwidpt 0.548 + 0.037 0.547 + 0.057 0. ai + 0,029 0.519 + 0.034 
(0.145) (0.171) (0.114) 097) 
(0.471; 0.616] (0.441; 0.612] (0. Hs 0.582] (0.473; 0.570] 
{0.535—0,.561} {0.522—0,572} {0.490—0.5 16} {0.500-0.538} 


flowering pedicel length and mean fruiting length are not significant in group 4 (P =0.15, 
F & 05 [1,20] = 2.23, F critical = 4.35) indicating that the inflorescence elongates very little 
from peak flower to mid-fruit. Significant differences between mean flowering and mean 
fruiting pedicel length were found in the other three groups. Group | has the largest F 
ratio (P < .OOOL, F «& .05 [1,44] = 20.39, F critical = 4.06) indicating that the lowest pedicel 
elongates significantly from peak flower to fruit. Group 2 and group 3 also show signifi- 
cant differences in this character (Group 2: P = .0O1, F « 95 [1.24] = 13.53, F eritical = 4.26; 
Group 3: P = .0006, F ».05[1.26)= 15.17, F critical = 4.23). 


DISCUSSION 


Multivariate analysis of morphological characters distinguishes four groups of specimens 
within the Amelanchier complex on the Delmarva Peninsula. Group | contains shrubby or 
multi-stemmed Amelanchier with short and rather fat petals, short, cylindrical flowering 
and fruiting inflorescences with short lower pedicels. This group of specimens is refer- 
able to Amelanchier obovalis; the type specimen nests within this group outside of the 
range of variation of other groups. This group may be more clearly visually defined by 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1041 


Taste 8. MANOVA test criteria and F approximations for the hypothesis of no overall effect of the four-group model. 
Statistic Value F Value Num DF Den DF Pr.F 

Wilk’s Lambda 0.04823940 14.12 18 150.39 ,_ 0001 

Pillai's Trace 147957567 7.29 18 165 , 0001 
Hotelling-Lawley 9.83905376 27.18 18 100.15 , 0001 


Taste 9. Univariate F values (GLM procedure) and P > F for six informative characters and pairwise group com- 
parisons of character means using Tukey’s Studentized Range (HSD). Critical value of Studentized Range = 3.74. 
*= Comparisons significant at a =.05, df =58; ns = not significant. Grp! = A. obovalis; Gro4 =A. canadensis. 


Character F ratio for P.F Group Difference Confidence Limits 
Character Comparison Ww Means 95% 

(Tukey) 

Flinfllen 61.59 0.0001 grp4-grp2 13.19 8.82—17.56 . 
grp4-grp3 8.25 3.95—12.55 . 
grp4-qrp1 19.10 15.21—22.98 * 
grp2-qrp3 4.98 0.827—9.05 
grp2-qrpl 5.9] 2.32—9.58 * 
grp3-grp| 10.85 7.26—14.43 * 

Flpedlen 68.86 ,0.0001 = grp4-grp2 3.70 2.01—5.39 * 
grp4-grp3 5.05 3.39—6.72 . 
grp4-grp1 7.93 42—9 
grp2-grp3 1.35 -0.24— 2.94 ns 
grp2-grp1 4.22 2.80—5.64 . 
grp3-grp1 2.87 1 48—4.26 . 

Flped_flinfl 10.16 ,0.0001 = grp4-qrp2 0.019 -0.032—0.070 ns 
grp4-grp3 0.063 0.012—0.113 . 
grp4-qrp] 0.053 0.007—0.098 - 
grp2-grp3 0.081 0.033—0.129 " 
grp2-grp| 0.071 0.028—0.114 i 
grp3-grp| 0.010 -0.032—0.052 ns 

Lfl_wratio 2.01 0.1228 grp4-grp2 0.104 -0.106—0.314 ns 
grp4-grp3 0.079 -0.127—0.285 ns 
grp4-grp| 0.016 -0.171—0.203 ns 
grp2-grp3 0.183 -0.015—0.380 ns 
grp2-grp | 0.088 -0.089—0.264 ns 
grp3-grp| 0.094 -0.078—0.267 ns 

Lfwidpt 5.99 0.0013 grp4-grp2 0.031 -0.009—0.072 ns 
grp4-grp-3 0.017 -0.023—0.057 ns 
grp4-grpl 0.030 -0.006—0.066 ns 
grp2-grp3 0.048 0.010—0.086 * 
grp2-grpl 0.002 -0.033—0.036 ns 
grp3-grp1 0.046 0.013—0.080 : 

Petlen 12.13 0.0001 grp4-grp2 1.11 0.079—2.15 ms 
grp4-grp3 1.53 0.510—2.55 [ 
grp4-grpl 2.06 1.14—2.98 : 
grp2-grp3 0.41 -0.559—1.39 ns 
grp2-grp1 0.94 0.073—1.81 * 


grp3-grp1 0.53 -0.320—1.38 ns 


1042 BRIT.ORG/SIDA 22(2) 


eliminating outliers; however, eliminating these individuals in the PCA does not change 
group membership and these specimens do appear to represent natural variation within 
the group. One individual classified in group 1 (upper right corner of Fig. 3) deviates from 
the morphological group defined here by an extremely narrow cylindrical inflorescence. 
This individual may best be recognized as a separate entity; however, no other similar 
individual or specimen was located during sampling. Individuals forming group | (A. 
obovalis) were collected chiefly from sandy woodlands and barrens but did occur on 
hummocks along tidal creeks and thicket margins around non-tidal wetlands in disturbed 
and in nearly pristine habitats. Individuals are typically low and shrubby but may grow to 
several meters in absence of disturbance; this was ideally illustrated by single, apparently 
clonal individuals with stems that were inside and outside of a roadside-mowing regime. 

Group 4 corresponds to a group of tall, arborescent or multi-stemmed Amelanchier 
with large petals and long lower pedicels forming long, widely spreading flowering and 
fruiting inflorescences. This group of specimens is referable to Amelanchier canadensis, 
matching as well as can be expected the type specimen and at least part of the historical 
circumscriptions. Individuals in group 4 (A. canadensis) were collected from both tidal 
and non-tidal swamp forests and were easily distinguished in the field. This species oc- 
curs sympatrically with Amelanchier obovalis in some habitats. 

Group land group 4 correspond to known taxonomic entities (Amelanchier obovalis 
and A. canadensis respectively) but group 2 and group 3, despite the statistical signifi- 
cance of several pairwise comparisons of character mean vectors in the MANOVA could 
not be assigned to distinctive forms such as might arise as microspecies. | hypothesize 
that group 2 and group 3 are members of an agamic complex derived from hybridization 
between Amelanchier canadensis and A. obovalis. Evidence of apomixis has been dem- 
onstrated in Amelanchier canadensis (Campbell et al. 1987) but not in A. obovalis. How- 
ever, evidence to support this hypothesis may be drawn from my statistical analysis and 
observation. First, both Amelanchier canadensis and A. obovalis (as clear entities) were 
observed to occur within the same habitat patch in the study area and were observed to 
have overlapping phenologies, which increase the probabilities that natural hybridiza- 
tion occurs. Second, the means of important classification characters in group 2 and group 
3 (with 90% confidence intervals) consistently occupy ranges intermediate between 
Amelanchier canadensisand A. obovalis (Table 7). Finally, the members of groups 2 and 3 
appear morphologically heterogeneous and do not correspond to the four putative mor- 
phological forms that I noted in the field prior to sampling or toa particular habitat type. 
My field observations were that one form was referable to Amelanchier obovalis(group 1),a 
second to Amelanchier canadensis (group 4),a third group comprised the pubescent-ovary 
forms within group | and the fourth, an early flowering Amelanchier with small petals on 

ummocks along tidal creeks | now believe are consistent with Amelanchier obovalis. 

Additionally, the discrimination of multivariate character space held by group 2 
versus group 3 is uncertain. Group 2 and 3 separate nearly at the origin of PCA-2 (Fig. 3) 
and the character most highly correlated with this second axis (r = .704) is Lfl_Wratio, a 
character that fails statistical significance tests between groups in Analysis of Variance 
(see Table 9). Further, comparison of mean vectors of two important characters (Flpedlen 
and Petlen) between group 2 and group 3 are not significant (Table 9). Finally, the itera- 
tions of Cluster Analysis join group 2 and 3 in a three-group model and then collapse 
into group 4 (A. canadensis) in a two-group model. If | accept more variation in 
Amelanchier canadensis and consider a two-group model composed of Amelanchier 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1043 


obovalis and a new group containing all the members of groups 2, 3 and 4 then my per- 
formance in keying vouchers would certainly improve but accepting this variation comes 
with a heuristic cost in that lobscure the only spatially distinctive cluster resulting from 
PCA. Given the well-documented propensity for hybridization and the formation of ag- 
amic complexes in Amelanchier, the most reasonable assumption is that individuals 
within group 2 and group 3 do constitute intermediate forms due to incomplete repro- 
ductive isolation between Amelanchier canadensis and A. obovalis. 

I conclude that the multivariate character space occupied by Amelanchier obovalis 
exhibits significant phenetic dissimilarities with the other groups as expressed by its 
distinctiveness in Cluster Analysis (e.g., the long branch before joining) and by statistical 
significance of the majorities of group comparisons of character mean vectors for 
Flinfllen, Flpedlen, Petlen, and Flped_flinfl. The discreteness of multivariate character 
space occupied by Amelanchier canadensis can be determined both visually (occurrence 
of a distinct cluster in the PCA) but also by the statistical significance of the majorities of 
group comparisons of character mean vectors for Flinfllen, Flpedlen, Petlen and 
Flped_flinfl. However, the presence of numerous intermediate specimens violates most 
commonly accepted taxonomic conceptions of species status. Subspecies as a taxonomic 
rank is poorly defined in the literature of taxonomic botany and is used more or less 
interchangeably with variety (McDade 1995; Nesom and Lipscomb 2005). Wiegand (1912), 
Fernald (1950a) and Jones (1946) assigned the rank of variety to those distinctive forms 
of Amelanchier that occupied a particular geographical area and/or had a habitat pref- 
erence slightly different than that typical of the species. Campbell and Dickinson (1990) 
concluded that nonconcordance between morphological and breeding units in Maloid 
apomicts rendered the biological speci ept inappropriate and that a morphospecies 
concept may be necessary to deal with evolutionary complexity in the Maloideae. In this 
study I found no well-defined geographical or ecological separation between the taxa. How- 
ever, I must also consider that the high frequency of intermediate forms found in the study 
area may reflect the long disturbance history of the Delmarva Peninsula and this distur- 
bance history is responsible for recent, but perhaps not historical, genetic and ecological 
exchangeability (Crandall et al. 2000). That the frequency of disturbance often correlates 
with the number of hybrids and ‘perplexing forms’ has been the subject of repeated obser- 
vation and much discussion among authors since Wiegand (1912). The study populations 
of Amelanchier fall under Crandall’s et al. (2000) case of “anthropogenic convergence on 
demographic exchangeability” where we continue to treat the populations as distinct units 
but not as distinct species. Both taxa are distinctive enough morphologically and interme- 
diate forms may not occur with aia frequency in other regions. In the taxonomic treat- 
ment below, I utilize the taxon ncept of “variety” defined as the primary taxonomic 
level below species ae Code of Botanical Nomenclature, 4.1). 


os 


TAXONOMIC TREATMENT 


Amelanchier canadensis (L.) Medic. var obovalis Michx. Typ spilu idensis var. obovalis 
Michx., Fl. Bor. Amer. 1:29]. 1803. Amelanchier obovalis (Michx.) pas Bor Gaz. 35:434. 1903, in part. 
Amelanchier obovalis (Michx.) Ashe, M.L. Fern ald in Rhodora 43:566. 1941, in part. Amelanchier 
obl bengvl ia (Torr. & A. Gray) ne K.M. Wiegand in yeaa 14:147-150. 1912, in part. Amelanchier 

obovalis, P Landry, Bull. Soc. Bot. Fr. 122:248. 1 


Amelanchier obovalis is a rather enigmatic species with a aie history of circumscrip- 
tion problems. André Michaux (1803) described Mespilus canadensis var obovalis from 


1044 BRIT.ORG/SIDA 22(2) 


“carolina inferiore” (lower Carolina) as “humilior; folius oblongiscule obovalibus” (low 
with oblong obovate leaves) and “arbriss de deux pieds haut’ (a shrub about two feet tall). 
The Michaux specimen (P) is very well preserved having several stems with inflorescences 
and petals intact and a single stem with (apparently mature) ded leaves. A century 
later Ashe (1903) created a new combination, Amelanchier obovalis (Michx.) Ashe de- 
scribing a shrub of “swamps and loose soils” 9-15 dm tall or a small tree to four meters 
tall with flowering inflorescences 3-5 cm long, the lowest pedicels 3-9 mm but becom- 
ing elongate (2-3 cm) in fruit. Wiegand (1912), probably the most influential of 
Amelanchier taxonomists, includes Amelanchier obovalis (Michx.) Ashe under his “fasti- 
giate not stoloniferous” Amelanchier oblongifolia Torr. & A. Gray) Roemer. stating that it 
“seemed to be this species” but it is not clear that Wiegand ever saw any of Ashe’s collec- 
tions from the southeastern United States. Wiegand (1912) described his A. oblongifolia 
as a shrub with slender upright stems in rather dense fastigiate clumps with inflores- 
cences 2.5-6 cm long, the lowest pedicels 8-18 mm long but scarcely longer in fruit. M.L. 
Fernald (1941), after several field seasons exploring the Pine Barrens of southeastern Vir- 
ginia, disagreed with Ashe’s description of Amelanchier obovalis, particularly regarding 
plant height, and subsequently ignored his treatment, describing Amelanchier obovalis 
(Michx.) Ashe as a “dwarf and stoloniferous” shrub with flowering inflorescences 1-2.5 
cm long with lower pedicels lengthening only to 3-8 mm. Fernald (1950a) distinguished 
the species in his key chiefly by plant habit in a major couplet: “stoloniferous or surculose 
loosely colonial shrubs” 0.2-2 M. high (A. obovalis); versus “fastigiate coarse and tall shrubs 
or arborescent” (A. canadensis). Jones (1946) closely followed Fernald’s (1941) treatment 
of Amelanchier obovalis as “low shrubs, surculose, forming loose colonies” but with flow- 
ering inflorescences 1-3 cm long with lower pedicels 1-3 mm, becoming 3-8(11-14 mm) 
long in fruit. Landry (1975) describes Amelanchier obovalisas a subspecies of A.canadensis 
with the major difference between the two subspecies being the general size of floral 
inflorescences and (very vaguely) the length of the lowest pedicels with Amelanchier 
canadensis ssp. canadensis having “rather large” flower clusters and lower pedicels at least 
6 mm long versus clusters of flowers small and lower pedicels more than 6 mm long (A. 
canadensis ssp. obovalis). 

The members of Group 1 (Amelanchier canadensis var. obovalis) are both low shrubs 
with apparent vegetative spread and tall, slender, multi-stemmed shrubs that occurred 
ina variety of habitats including hummocks on margins of tidal creeks and rivers, sandy 
pine forests and barrens, roadsides and under powerlines. In both flowering and fruiting 
stages the lowest pedicels were the shortest in the dataset and although they did elongate 
in fruit the extremes that Ashe (1903) suggested (10-20 mm) were not observed. I suspect 
that Ashe, perhaps due to finding very similar phenotypes in a variety of habitats or by 
witnessing the same complex as analyzed herein conflated the characteristics of both 
taxa. M.L. Fernald (1941), on the other hand, accepted only the extreme shrubby forms of 
the species and somewhat idiosyncratically noted that the taller species was probably 
Amelanchier dustromontana, a species that Ashe (1918) described from the Southern 
Appalachian Mountains and not the southern Coastal Plain. 


Amelanchier canadensis (L.) Medic.var. canadensis. Typr: Mespilus canadensis L., Sp. Pl. 1498. 1753. 


Amelanchier oblongifolia (Torr. & A. Gray) Roemer, K. Wiegand in Rhodora 14:147-150. 1912, in part. 
Amelanchier obovalis (Michx.) Ashe, Bot. Gaz. 35:434. 1903, in part. 


Amelanchier canadensis (L.) Medic. is the earliest combination in Amelanchier, the type 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1045 


specimen (LINN) from “habitat in Virginia, Canada” containsa single stem with five leaves 
and the remnants of a fruiting inflorescence with three fruit. The type is of limited use- 
fulness as the characters used by major authors of the genus to separate Amelanchier 
canadensis from A. obovalis, e.g. plant habit, petal length, flowering inflorescence length 
and lowest pedicel length are absent. Wiegand’s (1912) circumscription of Amelanchier 
oblongifolia appears to include both Amelanchier canadensis and A. obovalis as the ranges 
of character measurements are inclusive of both taxa. Similarly, Ashe’s (1903) circum- 
scription of Amelanchier obovalis appears to include cl istics of both species. It is 
very difficult to generalize but overall most authors treat Amelanchier canadensis asa 
fastigate shrub forming alder-like clumps having larger petals, and larger inflorescences 
than the low, shrubby and stoloniferous Amelanchier canadensis var. obovalis. 

The members of Group 4 (Amelanchier canadensis var. canadensis ) comprise a group 
of tall, fastigiate shrubs and small trees having the largest petal dimensions, very long 
flowering and fruiting inflorescences with long lower pedicels forming widely spread- 
ing inflorescences. This variety is easily distinguished in the field however numerous 
individuals with characters intermediate between the two varieties are present in the 
study area. Individuals of Amelanchier canadensis var. canadensis were located, mostly, 
in natural habitats associated with tidal and non-tidal wetlands, however, there appears 
to be little habitat specificity and these individuals occurred often in close proximity to 
var. obovalis. Wiegand’s notion that the pedicels of his Amelanchier oblongifolia are 
“scarcely longer in fruit” appears consistent with the results of Analysis of Variance com- 
paring lowest pedicel lengths in flowering versus fruiting material; although, it is appar- 
ent that Wiegand also conflated the cl teristics of both taxa, perhaps because he was 
unfamiliar with the more southern forms constituting Amelanchier obovalis. 


KEY 


The following key to Amelanchier canadensis includes the other common species occur 
ring in the study area, Amelanchier arborea (Michx. f.) Fernald. In keying Amelanchier 
one should realize that many specimens would not key confidently without observa- 
tions (collections) made throughout the growing season. Those with interme- 
diate characters are likely part of the species complex and so should be simply identified 
as ‘comparing favorably’ with one variety or the other. The twenty-four members of group 
1(A.c. obovalis) provide the mean and range (one standard deviation) of characters that 
should be used to identify this species and the eleven members of group 4 (A. c. canadensis) 
provide the mean and range of the same morphological characters. 

Specimens with entire leaves from the southeastern Coastal Plain may constitute 
var. subintegra Fernald but this putative taxon is not included in the key as its evaluation 
was not within the scope of this manuscript. 


Sepals in flower and in fruit tightly recurved and appressed to the ovary, leaves subcordate 7 
base, acute to acuminate at the apex and coarsely serrate. Arborescent species of ia 
renee nes arborea 
Sepals in flower and in fruit not tightly appressed to ovary, leaves rounded at base, ae to 
broadly rounded at apex and finely serrate. Tall shrubs with slender stems or dwarf shrubs with 
evident vegetative spread, chiefly of swamps and barrens at low elevations __ Amelanchier canadensis 


ioe (5.9-)6.8(-7.7) mm long and (2.6-)3.1(-3.6) mm wide, inflorescences at peak flower (27.8-) 
8.7(-29.6) mm, elongating in fruit to (27.2-)32.1(-37.1) mm, lowest pedicels (flowering) (6.6-) 

ah 9.2) mm, elongating in fruit to (8.2-)11.0(—13.8) mm. Inflorescences usually cylindrical, the 
lowest pedicel averaging 28% of the total inflorescence length at flowering, and 35% percent 


pee 


1046 BRIT.ORG/SIDA 22(2) 


in fruit. Leaves on older branches chiefly obovate, the widest point occurring at 51-58 percent 
of the total leaf length. Sepals usually erect in flower, but often spreading during maturation of 
the ovary.Ovary summit usually glabrous but a variable amount of pubescence occurs in some 
forms (typically in dry, well-drained habitats). Plants are typically short and shrubby but may 
w to several meters in the absence of disturbance var. obovalis 
ea (7.6-)8.8(-11) mm long and (3.1-)3.7(-4.3) mm ae inflorescences at peak flower (43.9-) 
47.8(-51.7) mm, elongating in fruit to (43.2-)48.2 (-53.2) mm, lowest pedicels (flowering) 
15.8(14.1)-(17.5) mm, elongating in fruit to (15.1-)17.5 (-19.9) mm. inflorescences using widely 
spreading, the lowest pedicel averaging 36% of the total inflorescence length at flowering, and 
38% in fruit. Leaves on older branches chiefly broadly elliptical or occasionally oblong, the wid- 
est point at 49-55% of the total leaf length. Sepals usually spreading to reflexed in flower and in 
fruit. Ovary summit glabrous. Plants are typically multi-stemmed with tall slender branches 
var. canadensis 


ACKNOWLEGMENTS 


[am grateful to Art Tucker, director, Robert Naczi, curator, and Keith Clancy, collections 
manager, of the Claude E. Phillips Herbarium (DOV), Delaware State Univ, Dover, DE for 
managing loans of herbarium material and providing access to a beautiful facility. Dur- 
ing the term of this research [have borrowed material from a number of institutions and 
I would like to thank the curators and staff at: CM, FLAS, GA, MARY, NA, NCU, PH, US, 
USCH, VPI, and WVA, who so efficiently handled the distribution of material to Dover. | 
would like to offer special thanks to Alain Changy (P) and Mark Spencer (LINN) who 
provided images of type specimens held in Paris and London, respectively. Gwen Brewer, 
Wesley Knapp and William McAvoy provided helpful comments on early drafts of the 
manuscript. Jason Harrison and Wesley Knapp assisted with fieldwork. Dan Sieh], with 
an artists care, mounted and processed hundreds of Amelanchier vouchers at the Tawes 
Herbarium, Maryland Department of Natural Resources, Annapolis, MD. The Maryland 
Department of Natural Resources, Natural Heritage Program, the Claude E. Phillips Her- 
barium, Delaware State University and the Norton-Brown Herbarium, University of Mary- 
land, College Park supported this research. Christopher Campbell, University of Maine, and 
an anonymous reviewer provided critical comments that greatly improved this manuscript. 


REFERENCES 

AsHe, W.W. 1903. Amelanchier obovalis. Bot. Gaz. (Crawfordsville) 35:434. 

AsHe, WW. 1918.Amelanchier austromontana. J. Elisha Mitchell Sci. Soc. 34:1 38. 

Baiey, R.G. 1995. Description of the ecoregions of the United States, Second Edition. USDA Forest 
Service, Washington, DC. Misc. Publ. No 1391 (rev.). 108 pp. with map at 1:7,500,000. 

Braun, E.L. 1950. Deciduous forests of Eastern North America. Blakiston Company, Philadelpia, PA. 

Brown, R.G.and M.L. Brown. 1972.Woody plants of Maryland. Port City Press, Baltimore, MD 

Campsett, C.S. 1999, The evolutionary role of hybridization in angiosperm agamic complexes, with 
special emphasis on Amelanchier (Rosaceae). In: Plant evolution in man-made habitats. L.W.D. 
Raamsdonk, and J.C.M. den Nijs, eds. Proceedings of the Vilth International IOPB symposium. 
Amsterdam, The Netherlands. Hugo de Vries Laboratory, Institute for Systematics and Population 
Biology, Univ. of Amsterdam, Pp. 341-358. 

Campsett, C.S.,C.W.Greene, and S.E.Berauist. 1987. Apomixis and lity in three species of Amelanchier, 
shadbush (Roasaceae, Maloideae). Amer. J. Bot. 74:321-328. 

Campsett, C.S., C.W. Greene, and TA. Dickinson. 1991, Reproductive biology in Subfam. Maloideae (Ro- 
saceae). Syst. Bot. 16:333-349. 


FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1047 


Campse.t, C.S., M.F. WosclecHowski, B.G. BALDWIN, L.A. Atice, and M.J. Donocuue. 1997b. Persistent nuclear 
ribosomal DNA sequence polymorphism in the Amelanchier (Rosaceae) agamic complex. Molec. 
Biol. Evol. 14:81-90. 

Campsett, C.S, and T.A. Dickinson. 1990. Apomixis, patterns of morphological variation, and species 
concepts in subfam. Maloideae (Rosaceae). Syst. Bot. 15:124-135. 

CampseLt, C.S. and W.A. WricHT. 1996. Apomixis, hybridization, and taxonomic complexity in Eastern 
North American Amelanchier (Rosaceae). Folia. Geobot. Phytotax. 31:345-354. 

Camesett, C.S.,W.A. WricHt, TF. Vininc, and W.A. Hatteman. 1997a. Morphological variation in sexual and 
agamospermous Amelanchier (Rosaceae). Canad. J. Bot. 75:1166-1173. 

CRANDALL, K.A., O.R.P. BininpA-Emonps, G.M. Mace, and R.K. Wayne. 2000. Considering evolutionary pro- 
cesses in conservation biology. Trends Ecol. Evol. 15:290-295. 

Cruise, J.E. 1964. Studies of natural hybrids in Amelanchier. Canad. J. Bot.42:65 1-663. 

Disete, A.C.,W.A. WRIGHT, C.S. CampBeLt, and C.W. Greene. 1998. Quantitative morphology of the Amelanchier 
agamic complex (Rosaceae) at a Maine site. Syst. Bot. 23:31- 41. 

Dickinson, T.A.and C.S.Campsett. 1991.Population structure and reproductive ecology in the Maloideae 
(Rosaceae). Syst. Bot. 16:350-362. 

Fernato, M.L. 1941. Another century of additions to the flora of Virginia. Rhodora 43:559-630. 

Fernaco, M.L. 1946. Amelanchier spicata not an American species. Rhodora 48:125-134. 

Fernaco, M.L. 1950a. Gray’s manual of botany, 8th edition. Dioscorides Press, Portland, OR. 

Fernato, M.L.1950b. A southern Amelanchier. Rnodora 52:67-68. 

Geason, H.A.and A. Cronauist. 1991. Manual of the vascular plants of northeastern United States and 
adjacent Canada. New York Botanical Garden, Bronx. 

Houmaren, P.K., N.H. Houmeren, and L.C. Barrett. 1990. Index herbariorum. Part 1.Edition 8.Regnum Veg. 
120:x, 1-693. 

Jackson, D.A. 1993. Stopping rules in principal components analysis:a comparison of heuristical and 
Statistical approaches. Ecology 74:2204-2214. 

Jones, G.N. 1946.American species of Amelanchier. Illinois Biol. Monogr. 20(2) 

Lanory,P.1975.Le concept _espéce et la taxinomie du genre Amelanchier (Rosacées). Bull. Soc. Bot. Fr. 
122:243-252. 

Leonaro, M.LR., R.E. Cook, and J.C. Sempce. 2005. A multivariate morphometric study of the Aster genus 
Sericocarpus (Asteraceae: Astereae). Sida 21:1471-1505. 

Linnaeus, C. 1753. Species Plantarum. Vol. 1. 

McCune, B.and J.B. Grace. 2002. Analysis of ecological communities. MjJM Software Design. Gleneden 
Beach, OR. 

McCune, B. and M.J. Merrord. 1999. Multivariate Analysis of Ecological Data, Version 4.14. MjM Soft- 
ware, Gleneden Beach, Oregon, USA. 

McDaoe, L.A. Species concepts and problems in practice: insight from botanical monographs. Syst 
Bot. 20:606-622. 

Michaux A. 1803.Flora Boreali-Americana, In:J.Ewan,ed.Classica Botanica Americana, Volume 3.Hafner 


Press, New York. 

Nesom, G.L. and B. Lipscoms. 2005. ICBN clarification needed: use of ranks. Sida 21:2185-2191. 

Nietsen, E.L. 1939. A taxonomic study of the genus Amelanchier in Minnesota. American Midland 
Naturalist 22:160-208. 

Puipes, J.B., K.R. Rogertson, PG. Smit, and J.R. Ronrer. 1990. A checklist of the subfamily Maloidae (Ro- 
saceae). Canad. J. Bot. 68:2209-2269. 

Raproro, A.E.,H.E.Ahles,and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. Univ. of North 
Carolina Press, Chapel Hill. 


1048 BRIT.ORG/SIDA 22(2) 


Rogertson, K.R. 1974. The genera of the Rosaceae in the southeastern United States. J. Arnold Arbor. 
55:633-662. 

RoserTson, K.R., J.B. Pripes, and J.R. Ronrer. 1992. Summary of leaves in the genera of Maloideae (Ro- 
saceae). Ann. Missouri Bot. Gard. 79:81-94. 

Rosinson, W.A. 1982. Experimental in the genus Amelanchier. ll. Do the taxa in Amelanchier form an 
agamic complex? Rhodora 84:85-100. 

Rosinson, W.A. 1986. Effect of fruit ingestion on Amelanchier seed germination. Bull. Torrey Bot. Club 
113:131-134. 

Saareta, J.M., BM. Peterson, RJ. Sogenc, and R.E. CHapman. 2003. A taxonomic revision of the Eastern 
North American and Eastern Asian disjunct genus Brachyelytrum (Poaceae): evidence from mor- 
phology, phytogeography and AFLP'’s. Syst. Bot. 28:674-692. 

ScHmibt, M.F., Jk. 1993. Maryland's geology. Tidewater Publishers, Centreville, MD. 

Sokat, R.R.and F.J. Ronit. 1969. Biometry. W.H. Freeman and Company, San Francisco, CA. 

Tatnall, R.R. 1946. Flora of Delaware and the Eastern Shore. Society of Natural History of Delaware, 
Lancaster, PA. 

Weser, J.E. and C.S. Camesett. 1989. Breeding system of a hybrid between a sexual and an apomictic 
species of Amelanchier, Shadbush (Rosaceae, Maloideae), Amer. J. Bot. 76:341-347. 

WHANG, SUNG S00, KYUNG CHOI, R.S. Hitt, and JAe-HOoNG Pak. 2002.A morphometric analysis of infraspecific 
taxa within the /xeris chinensis complex (Asteraceae, Lactuaceae). Bot. Bull. Acad. Sin. 43:131-138. 

Wiecand, K.M.1912.The genus Amelanchier in eastern North America. Rhodora 14:117-161. 

Wiecanb, K.M. 1920. Additional notes on Amelanchier. Rhodora 22:146-151. 

WitcaNb, K.M. 1935.A taxonomist's experience with hybrids in the wild. Science 81:161-166. 


GAREX REZNICERILA NEW WIDESPREAD SPECIESOF 
CAREX SECTION ACROCYSTIS (CYPERACEAE) 
FROM EASTERN NORTH AMERICA 


David A.Werier 


30 Banks Road 
Brooktondale, New York 14817, U.S.A. 
nakita@lightlink.com 


ABSTRACT 


Carex reznicekii is described as new from mesic to dry-mesic forests of Alabama, Arkansas, Connecticut, Dela- 
ware, District of Columbia, Georgia, Kentucky, Maryland, Mississippi, Missouri, New Jersey, New York, North 
Carolina, Pennsylvania, Rhode Island, South Carolina, Tennessee, and Went pees C. reznicekii had mostly 
been misidentified as either C. umbellata or C. nigromarginata. It is distingui d from these species as well as 
other members of section Acrocystis by a combination of lack of pasa) spikes, once culms, narrow leaves, and 
only ce red pistillate scales. A key to the 19 taxa of section Ac t rth America is 
provided. 


RESUMEN 


Se d ibe como nuevo Carex reznicekii de los bosques mésicos a xérico-mésicos de Alabama, Arkansas, Caro- 
lina del Norte, Carolina del Sur, Connecticut, Delaware, Distrito de ae Georgia, Kentucky, Maryland, Mis- 
sissippi, Missouri, New Jersey, New York, epee ibis Rhode Island, Tennessee, y Virginia. Previamente, C. 
bee habia ace en su mayor parte mal interpretada 3G, aan lataoC. eee a. Se distingue 
o de otros miembros de ie seccion eee ae por la combin cin de carencia de espiga 
basal, nee cor tos, ee Soa cn2s) jas. Se proporciona una clave para 
los 19 taxa de la seccion Acr s que ocurren en el este de Norteamérica. 


INTRODUCTION 


Carex section Acrocystis Dumort. contains approximately 35 species worldwide with 20 
in North America (Crins & Rettig 2002), although the most recent comprehensive treat- 
ment is Kikenthal’s (1909) now outdated revision. Species distributions lie mostly in 
Eurasia and North American with one taxon known from South America (Ktkenthal 
1909). It appears that Carex section Acrocystis as currently circumscribed is polyphyl- 
etic, but North American species and some Eurasian species form a clade (Roalson et al. 
2001, Roalson & Friar 2004). Twenty-eight taxa, including the one described here, are 
currently known from North America, with 19 of these occurring in eastern North 
America (Crins & Rettig 2002). At least two additional undescribed species occur in east- 
ern North America (Crins & Rettig 2002; Roalson & Friar 2004; Anton Reznicek, pers. 
comm). 

Eastern North American taxa of section Acrocystis, share the following characters: 
perigynia pubescent (except C. tonsa (Fernald) Bicknell var. tonsa ), two veined, and 
abruptly narrowed toa distinct beak; stigmas three (except two and three in C.floridana 
Schwein.); non-basal pistillate spikes (when present) subtended by sub-sheathing to 
sheathless bracts, approximate, the distal ones short pedunculate to sessile; and habitats 
dry to mesic forests (tending to favor drier) or more open for a few taxa. 

During survey work in the Hudson Highlands of southeastern New York, I found a 


SIDA 22(2): 1049- 1070. 2006 


1050 BRIT.ORG/SIDA 22(2) 


population of the New York State endangered C. nigromarginata. While delineating and 
describing this population I encountered a population of a strikingly different plant, 
which at first appeared to be C. umbellata because all of its culms were very short and 
hidden in the bases of the plants. Upon closer inspection, | observed that these short- 
culmed plants lacked the basal spikes characteristic of C. umbellata. These short-culmed 
plants were uniform throughout its population and appeared immediately adjacent to 
and clearly did not resemble C. nigromarginata. In contrast to the C. nigromarginata popu- 

ation, these short-culmed plants had much narrower leaves, uniformly very short culms, 
and pale pistillate scales. Wider sampling showed that the short-culmed plants formed 
discreet uniform populations, although sometimes occurring mixed with, but seemingly 
not intergrading with, C. nigromarginata. A cursory examination of herbarium material 
also suggested that the distributions of the two entities were different. This new species 
of Carex is described here as Carex reznicekii. 


MATERIALS AND METHODS 


Specimens of C. reznicekii were sought by examining all specimens labeled as C. 
nigromarginata Schwein. and C. umbellata Schkuhr ex Willd. (two superficially similar 
species) and in some cases other members of section Acrocystis from BH, BRIT, GH, MO, 
NCU, NY, NYS, PH, and US. In addition, putative C. reznicekii specimens were sent to me 
from collections at DOV, MICH, and VPI as well as by Philip E. Hyatt. Additional speci- 
mens were gathered from field work. Type material was examined of all species similar 
to or confusable with C. reznicekii, including their synonyms. 

Thirty-one populations were visited in the field, covering a large part of the species’ 
range. Site visits were conducted to assess habitat affinities, morphological variability, 
and habitat differences with other members of Carex section Acrocystis. Soil samples were 
collected at six distant sites throughout a large part of the range of the species (Fig. 1). 
Equal amounts of soil from four places at each of the six sites (adjacent to individual C. 
reznicekii plants) that were at least 10 meters apart were mixed together. These were sent 
to Cornell Nutrient Analysis Laboratories at Cornell University, Ithaca, NY for analysis. 
Soils were analyzed for particle size distribution, pH, and the minerals Al, Ca, Fe, K, Mg, 
Mn, P and Zn. 

To clarify the differences between C. nigromarginata, the most similar species to C. 
reznicekii,and C. reznicekii, specimens were compared using eight continuous charac- 
ters (see Table 1). Specimens were selected so as to have ten specimens per species from 
each of three geographic regions: 1) northeast (Virginia/Kentucky north), 2) southeast 
(North Carolina/Tennessee south), and 3) Arkansas/Missouri. Only six adequate speci- 
mens were available for C. reznicekii from the latter region. Within each geographic re- 
gion (excluding Arkansas/Missouri), only two specimens per state and one per county 
were selected to help cover the range of the species. For each species, specimens were 
selected randomly given the r ioned above and excluding specimens where 
all eight characters could not be measured. Analysis of variance (two-way ANOVA) and 
Pearson Correlation Coefficients were calculated using SPSS Version 13. Additionally, 
ranges, means, and standard deviations were calculated for both species for each region. 

Nonmetric multidimensional scaling (NMDS) ordination was chosen to compare 
the relationship of C. nigromarginata and C. reznicekii. In comparison to other ordina- 
tion methods such as canonical correspondence analysis (CCA) or principle components 
analysis (PCA), NMDS does not assume an underlying distribution for measured vari- 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1051 


pares 


| ee ay: 


= ATE BLT racae es 
Tay ines Hh apaue zine 


en 


oy 
Serene cee ieee ro Z = 


La TS 


a a" on 


aeness 
Lt 


—+ 
iT 


Cail I Hartad in hla-l 1 rs 


Fic. 1.G hic distributi f C. reznicekii 
Gla} 


ables. A NMDS ordination was created using the slow thorough mode on PC-ORD 
(McCune & Grace 2002). Specimens measured for use in the ordination are delineated by 
a single asterisk in the isotypes listed for C. reznicekii, in the representative specimens of 
C. reznicekii, and in the specimens cited in Appendix A. 

A scatter plot graph using a larger sample size was created for the two most explana- 
tory characters. As additional evidence that C. reznicekii and C. nigromarginata are dis- 
creet entities, distribution curves were created and compared for the two most explana- 
tory characters for C. nigromarginata and C. reznicekii, as well as C. nigromarginata and 
C. reznicekii combined, and C. nigromarginata specimens from outside the geographic 
range of C. reznicehii. For the latter distribution curves all sp of C.nigromarginata 
that occur outside of the geographic range of C. reznicekii from BH, BRIT, MICH, and 
MO, excluding duplicates, were measured. Specimens measured for the scatter plot graph 
and distribution curves are delineated by either a single or double asterisk in the isotypes 
listed for C. reznicehii, in the representative specimen of C. reznicehii, and in the speci- 
mens cited in Appendix A. 

description of C. reznicekii and a key to all members of Carex section Acrocystis 
were created from original measurements except, for the key, measurements for C. com- 
munis Bailey varieties and C. inops Bailey ssp. heliophila (Mackenzie) Crins are adapted 
from Crins and Rettig (2002). Character states that were on an extreme edge or disjunct 
from most other measurementare placed in parentheses. The following notes clarify three 
characters listed in the description and the key. 

1. Culm length is measured from the base of the culm to the top of the inflorescence 
as interpreted or implied by many other authors (LeBlond et al. 1994; Reznicek & 
Camelbeke 1996; Naczi et al. 2001; and others). 


2. Terminal staminate spikes exceeding or exceeded by the rest of the inflorescence is 


1052 BRIT.ORG/SIDA 22(2) 


Taste 1. Characters measured for ordination, ANOVA, and Pearson Correlations giving the codes used through- 
out the paper and any specifications for choosing which part to measure. 


Character Code Specifications 

Tallest culm height (cm) TaCuHe Tallest per specimen 

Widest leaf width (mm) ea Widest per specimen 

Anther length (mm) ean of two per specimen 
Largest length that a staminate setae Greatest length per specimen 


spike exceeds the dista 
most pistillate spike (mm) 


Perigynia length (mm) PeLe Mean of two per specimen 

Perigynia width (mm) PeWi Mean of two per specimen 

Taller staminate spike length (mm) StSpLe Mean of the length of two taller staminate 
spikes per specimen 

Staminate spike width (mm) StSpWi Mean of the width of the two staminate spikes 


measured for spike length 


measured by subtracting the difference of the length from the base of the bract subtend- 
ing the proximal-most non-basal spike to the apex of the terminal staminate spike, and 
the ae from the base of this same bract to the apex of the distal most lateral spike. 

3. Perigynium body refers to the perigynium excluding the beak and stipe or stipe- 
like base. The beak and stipe begin at the deepest point in the concavity formed at the 
summit and base of the perigynium. Perigynium body shapes vary considerably and al- 
though their shape aids in distinguishing taxa (and is used in the key), they should be 
used cautiously. 


RESULTS 
Carex reznicekii Werier, sp. nov. (Fig. 2). Typr: U.S.A. VirGiNiA. Caroline Co: near North Anna River, 
Route 207 Jericho Road) W from Carmel Church to Oxtord Road (Route 689), 3.6 mi S and SE on Oxford 
Road (Route 689) to small dirt road on S side of road; SW along dirt road towards North Anna River, UTM 
coordinates in NAD-83 Zone 18 4196701N 280996E, 10 Apr 2004, Werier 1951 (HOLOTYPE: BH; ISOTYPES: 
GH*, MICH, MO, NCU, NY, US). 
Carici nigromarginatae similis, sed characteribus sequentibus differt: laminis foliorum latissimis 1.2-2.2(-2.5) 
mm latis; culmis singulae plantae saepe in longitudine similibus longissimis 1.9-9.9(-13.7) cm pe spicis 
staminalibus 3.3-8.3(-9.2) mm longis et 0.4-1.6(-1.8) mm latis; et colore rubenti squamarum pistillatarum ad 
regiones marginales restricto. 
Densely caespitose, herbaceous, evergreen perennial. Rhizomes horizontal, ascending, or 
erect, 0.1-3.8 mm long between shoots or branches of the rhizomes, 0.6-1.2 mm wide with 
leaf sheaths absent, enclosed by leaf sheaths and/or cataphylls, which disintegrate into 
long fibers. Vegetative shoots erect to ascending, arising from buds at the tip or side of the 
rhizomes, or directly from the apical meristem of rhizomes or previous year's vegetative 
shoots; bases consisting of cataphylls or leaf sheathes which disintegrate into long fi- 
bers; pseudoculms (represented by leaf sheaths) 1.2-7.6 cm long. Reproductive shoots erect, 
mostly produced directly from the apical meristem of vegetative shoots or rhizomes, or 
less frequently produced from buds at tip or side of rhizomes, bases mostly surrounded 
only by leaf sheaths which disintegrate into long fibers, or less frequently surrounded 
by cataphylls; culms 1.3-9.9(-13.7) cm long, the tallest 1.9-9.9(-13.7) cm long, 0.3-0.5 mm 
wide just below the inflorescence, trigonous, with 1 elevated vein on each angle and 2-5 
elevated veins on each side, smooth to minutely antrorsely scabrous on angles and veins, 


1053 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 


1mm).Middle 


Fig. 2. Carex reznicekii. Bottom right, plant (scale 
(I to r), culm, inflorescence, and sheath (scale = 1 cm). Top left, staminate scale and anther (scale = 1 mm). 


, perigynia, and achenes (scale 


ft, Pistillate scales 


a 


5 cm).Bottom le 


1054 BRIT.ORG/SIDA 22(2) 


more prominently scabrous on angles and distally. Cataphylls at base of reproductive 
shoots 0-2, 1.6-10.8 mm long. Leaves of reproductive shoots 3-9. Leaf blades to 52.0 cm 
long, 0.7-2.2(-2.5) mm wide, the widest 1.2-2.2(-2.5) mm wide, flat to V-shaped, to occa- 
sionally M-shaped; adaxial surface dark green and smooth, papillose, or antrorsely sca- 
brous, more textured distally; abaxial surface smooth to occasionally scabrous, more tex- 
tured on mid-vein and distally; margins smooth to antrorsely scabrous, more scabrous 
distally; leaf sheaths 0.7-5.1 cm long, lower portion especially on outer sheaths have some 
red coloration; abaxial face with green to white or proximally red veins and whitish and 
translucent to proximally reddish and more opaque intervein regions; adaxial face with 
whitish or proximally red veins and whitish, thin, and translucent intervein regions, thin- 
ner and more translucent than abaxial intervein regions; vein edges with ascending, 
spreading, or reflexed minute, stiff, broad-based deltoid hairs; sheaths disintegrate into 
long fibers consisting of veins, which retain pubescence; adaxial sheath face apex con- 
cave to a depth of 0.3-0.8(-1.0) mm and sometimes slightly thickened; ligules typically 
wider than long, 0.3-1.0(-1.5) mm long (including the free portion), 0.6-1.4 mm wide, free 
portion of ligule 0.1-0.2 mm long, ciliate with abundant minute, stiff, broad-based del- 
toid hairs. Infructescences 5.2-12.9(-16.7) mm long, consisting of 3-5 approximate spikes 
at the summit of the culm; proximal-most internode 0.8-3.7(-5.2) mm long; bracts 
sheathless; proximal-most bract with blade 3.6-23.0(-38.5) mm long, 0.6-1.8 mm wide, 
3.9(-4.3) mm shorter than to 10.9(-23.5) mm taller than infructescences; distal bracts re- 
duced. Spikes arising singly from nodes; terminal spike staminate, 3.3-8.3(-9.2) mm long, 
0.4-1.6(-1.8) mm wide, on peduncle (0.2-)0.3-0.7 mm long, exceeding distal-most lateral 
spike by 3.7 mm to exceeded by distal-most lateral spike by 1.1(-1.6) mm; lateral spikes 2- 
4, pistillate, short-pedunculate, with a cladoprophyll towards base of peduncle; proxi- 
mal-most spikes 3.4-7.2 mm long, (1.8-)2.2-4.4 mm wide, (3-)5-10(-12) flowered, on pe- 
duncles (0.3-)0.4-0.9(-1.4) mm long, with cladoprophyll (0.7-)1.7-2.8(-3.0) mm long. 
Staminate scales (2.5-)2.7-4.0 mm long, 0.7-1.8 mm wide, ovate to elliptic, apex acute to 
obtuse, with a green or yellowish longitudinal mid-stripe which includes the mid-vein, 
margins translucent, thin, whitish to occasionally yellowish-brown, often red sub-mar- 
ginally forming a longitudinal stripe, more so toward apex of scales and on distal scales, 
mid-vein present, extending further toward the apex of the scale in more distal scales; 
distal scales with mid-vein raised, slightly antrorsely scabrous, and ending close to apex of 
scale or sometimes with mid-vein projecting as a short awn attached just proximal to the 
scale apex and projecting up to just beyond the apex of the scale; Pistillate scales 2.6-4.3 
mm long, 1.0-1.9 mm wide, 0.9(-1.) mm shorter than to 0.5 mm longer than associated 
perigynium, ovate to lanceolate to elliptic, apex acute to obtuse, with a green (to brownish) 
longitudinal mid-stripe which includes the mid-vein; margins translucent, thin, whitish 
to occasionally yellow-brown, often some reddish color sub-marginally forming a sub- 
marginal longitudinal stripe, more so distally on the scale; sub-marginal longitudinal red 
stripe Gif present) does not extend laterally to immediately adjacent to mid-stripe except 
sometimes at apex of scale; mid-vein antrorsely scabrous and ending just before apex of 
scale or mid-vein, projecting as a short awn attached just proximal to the scale apex and 
projecting up to just beyond the apex of the scale; bases of scales often with a narrow red 
horizontal stripe just above attachment to spike axis. Anthers 3, (1.2-)1.3-1L9(-2.3) mm long. 
Stigmas 3, withering a age. Perigynia (2.5-)2.7-3.9 mm long, (0.8-)0.9-1.3(-1.5) mm wide, 
light green sparsely red-punctate, obtusely trigonous to plano-convex in cross- 
section, with two prominent nerves that extend the length of the perigynia and occasion- 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1055 


ally up to8 less nerves at base, papillose with papillae short-cylindric up to 0.01 
mm. long, as well as pubescent with small (less than 0.1 mm), predominately antrorsely 
directed, stiff, deltoid hairs, which are denser on the prominent nerves that extend into the 
beak teeth; body 1.4-1.9(-2.0) mm long, ellipsoid, gradually tapering to beak and stipe: stipe 
0.5-1L0(-LD mm long; beak (0.4-)0.6-1.0 mm long, bidentate; beak teeth 0.1-0.3 mm long. 
Achenes (1.4-)1.5-18(-19) mm long, 0.9-1.2(-1.3) mm wide, ovoid, acutely to obtusely 
trigonous to plano-convex in cross-section, minutely papillose, yellow-green to green when 
immature, light to chestnut brown when mature. 

Etymology.—I selected reznicekii as the epithet to honor Dr. Anton Reznicek, Univer- 
sity of Michigan, who has inspired me and many others interested in carices. He has cata- 
lyzed a revival in the study of Carex which has revealed a tremendous amount of new 
information. In addition, he has contributed directly toa greater understanding of these 
plants through numerous articles and classes. 

A total of 242 specimens (143 records) of C. reznicekii were examined. Of these, 125 
specimens (88 records) were collections made prior to recognition of C. reznicekii. Sev- 
enty-one of these 125 specimens had at one point been labeled C. nigromarginata, 78 C. 
umbellata (including C. abdita Bicknell and C. umbellata var. brevirostris Boott), 6 C. 
floridana [including C. nigromarginata var. floridana (Schwein.) Ktikenthal], and 9 C. 
albicans Willd. ex Spreng. var. emmonsii (Dewey ex Torr.) Rettig [including C. emmonsii 
Dewey ex Torr.and C. nigromarginata var. minor (Boott) Gleason]. The sum totals to more 
than 125 because some specimens had been annotated numerous times. 

Type material examined comprised the following: C. umbellata, scan of holotype from 
B; C. umbellata var. vicina Dewey, holotype at GH, isotype at PH; C. umbellata var. 
brevirostris, isotype at US (holotype not examined but specimen is from Saskatchewan): 
C. abdita, holotype at NY; C. microrhyncha Mackenzie, holotype at NY: C. umbellata var. 
tonsa Fernald, lectotype and 2 isolectotypes at GH; C. rugosperma Mackenzie, holotype at 
NY, C. nigromarginata, holotype at PH; and C.floridana, holotype at PH. None of the types 
examined proved to be C. reznicekii. 

A detailed comparison of C. reznicekii to C. umbellata and C. nigromarginata is made 
below. In the past, C. reznicekii has been mistaken most often for these two species. Carex 
nigromarginata is most similar. The key compares all members of section Acrocystis from 
eastern North America. 

Carex umbellata superticially resembles C. reznicekii, but is actually quite distinct. 
The two species can easily be fully separated by noting the presence (C. umbellata) or 
absence (C. reznicekii) of basal spikes. Mackenzie (1913), clearly stated this when he ex- 
plained how to distinguish C. nigromarginata and C_floridana from C. umbellata and its 
allies. He stated, “these [C. nigromarginata and C.floridanal]are to be distinguished by the 
fact that while the spikes are on very short culms and may appear basal they are not on 
basal peduncles.” Fernald (1902) apparently did not understand the concept of basal 
spikes when he stated that “the best means of distinction between C. umbellata [misap- 
plied to C. tonsa var. rugosperma (Mackenzie) Crins] and C. nigro-marginata [sic] is of- 
fered by the thickness of the perigynia.” In his 8th edition of Gray’s Manual, Fernald (1950) 
placed C. nigromarginata with the basal spike members of section Acrocystis (as Montanae 
Fries) and then used perigynium characters and geographic region to distinguish C. 
nigromarginata from C. umbellata and its allies. This means of distinction appears to have 
been followed by at least Cusick (1992) and Tucker (1995), and may have in part resulted 
in C. reznicekii often being misidentified as C. umbellata. 


1056 BRIT.ORG/SIDA 22(2) 


Other characters that separate C. umbellata from C. reznicekii include the following: 
C. umbellata has staminate spikes with 0-2 approximate pistillate spikes vs. 2-4 for 
reznicekii; some pistillate spikes on elongate peduncles vs. none on elongate peduncles 
for C. reznicekii; staminate spikes extending up to 9.0 mm beyond the approximate (if 
present) pistillate spikes vs. staminate spikes at most extending 3.7 mm beyond the pis- 
tillate spikes for C. reznicekii; and perigynia (2.2-)2.3-3.2(-3.3) mm long vs. perigynia 
(2.5-)2.7-3.9 mm long for C. reznicekii. In addition, leaf blades of C. umbellata are gener- 
ally lighter green and slightly wider than C. reznicehii. 

A full understanding of basal spikes seems particularly important to understand- 
ing . difference between these two species, and may enrich the understanding of sec- 
tion Acrocystis as a whole. Basal spikes are individual pistillate spikes which arise al- 
most directly from the base of the plant [subradical of Mackenzie (1913)]. These pistillate 
spikes have elongated peduncles, and as with all pistillate spikes in section Acrocystis, 
they are subtended by bracts. These bracts have sheaths and blades [contrary to Macken- 
zie (1935) and Roalson and Friar (2004) that state that section Acrocystis (Montanae of 
Mackenzie) has bracts which are sheathless or subsheathing]. These authors probably 
meant that bracts of non-basal spikes are sheathless or subsheathing. In addition, the 
culms of C. umbellata (defined as from the bases of the culms to the apices of the inflo- 
rescences) are often much taller than the apices of the basal pistillate spikes. Roalson 
and Friar (2004) consider basal pistillate spikes to be better described as a continuous 
character with somewhat distinct states, as opposed to a discrete character. While in es- 
sence this could be true, it appears that this character functions in two completely dis- 
tinct states with associated other characters (elongated peduncles and sheathing bracts). 
Even more interestingly, while neither C. nigromarginata nor C. reznicekii has basal 
spikes, both of ten have “grouped culms.” These “grouped culms” have one culm which is 
taller and one to three which are shorter (much less pronounced in C. reznicekii), mim- 
icking the tall and short aspect of C. umbellata, with its often tall culms and shorter basal 
pistillate spike peduncles. In addition, the shorter culms in a “group” of culms in C. 
nigromarginata and C. reznicekii have relatively shorter and less projecting terminal 


staminate spikes. 

Carex reznicekii and C. nigromarginata share a few characters, including no basal 
spikes, no elongated rhizomes, at least some culms short and hidden in leaf bases, 
perigynia of similar length, stigmas 3, and inflorescences composed of 3-5 approximate 
spikes. Likely because of these similarities, C. reznicekii has often been misidentified as 
C. nigromarginata. 

Several characters separate these two species. First, C. reznicekii has narrower leaves 
with the widest per plant 1.2-2.2(-2.5) mm wide vs. (1.9-)2.3-4.5 mm wide for C. 
nigromarginata. Second, the tallest culms per plant are shorter. For C. reznicekii the tall- 
est culms are 1.9-9.9(-13.7)cm long vs. (4.5-)6.6-38.0(-51.0) cm long for C. nigromarginata. 
Third, individual plants of C. reznicekii have culms often about the same length com- 
pared to widely different lengths among individuals of C. nigromarginata. Fourth, C. 
reznicekii culms remain erect even at maturity while some culms droop at maturity for 
C. nigromarginata. Fifth, C. reznicekii has margins of pistillate scales without red or red 
only submarginally. The red coloration (if present) can form a submarginal longitudinal 
stripe, but does not extend laterally to the green (or brown) longitudinal mid-stripe of 
the scale except at the apex. Carex nigromarginata has pistillate scales either similar in 
color to C. reznicekii or more often with dark reddish to dark purplish/black coloration 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1057 


that extends laterally from the submargin to the green (or brown) mid-stripe of the scale. 
In addition, the red to purple color often extends all the way to the base of the scale in C. 
nigromarginata, while in C. reznicekii the reddish color (if present) does not extend to 
the base of the scale. Sixth, the apex of the staminate spike in C. reznicekii exceeds the 
apex of the distal-most lateral spike by at most 0.0-3.7 mm. In C. nigromarginata, the 
apex of the staminate spike which exceeds the apex of the distal-most lateral spike the 
most, exceeds it by (0.0-)0.5-5.\(-7.3) mm. While this character exhibits considerable 
overlap in these two species, it can sometimes be useful for distinguishing difficult speci- 
mens. Seventh, C. reznicekii has staminate spikes on average slightly shorter and nar- 
rower, measuring 3.3-8.3(-9.2) x 0.4-L.6(-18) mm vs. 4.2-10.9(-12.0) x 0.5-2.4(-2.8) mm 
for C. nigromarginata. Again this character shows significant overlap, but it occasionally 
provides useful help in distinguishing between these two species. 

Carex reznicekii and C. nigromarginata were compared for eight continuous charac- 
ters (see Table 1). Ranges, means, and standard deviations for all eight continuous charac- 
ters measured for the NMDS ordination are presented in Table 2. Comparing C. 
nigromarginataand C. reznicekii, PC-ORD recommended a two-dimensional solution for 
the NMDS ordination. This resulted in a final ordination (see Fig. 3) with low instability 
(<0.001) indicating convergence of the iterations, and low stress (6.61) indicating that the 
data was far from random and there was “no real risk of drawing false inferences” (McCune 
S& Grace 2002). Axis 1 of the ordination represents decreasing TaCuHe (-0.961) and 
WiLe Wi (-0.747) with the numbers in parentheses representing the Pearson Correlation 
Coefficients with Axis 1 (Fig. 3). Axis 1 explained 81% of the variability in the dataset. 
Axis 2 of the ordination represents increasing TaCuHe (0.772), WiLeWi (0.707), 
StSpExPiSp (0.883), StSpHe (0.899), and StSpWi (0.702) with the numbers in parentheses 
representing the Pearson Correlation Coefficients with Axis 2 (Fig. 3). Axis 2 explained 
an additional 17% of the variability for a total of 98% explained variability. Overall, the 
ordination summarizes a very Se dataset with several highly correlated variables 
and in addition shows that while the two species are distinct, some overlap of individual 
specimens exists. Pearson-correlations illustrated that all of the characters measured 
except width and length of the perigynia were strongly correlated with one another (p< 
0.01), indicating a high level of redundancy of the characters used. The ordination is pre- 
sented mainly as a visual tool to show how C. reznicekii and C. nigromarginata compare 
with numerous characters. 

All of the characters measured except width and length of the perigynia differed 
significantly between C. reznicekii and C. nigromarginata (Table 3). The scatter plot graph, 
using a larger sample size for the two strongest characters separating the two species (tall- 
est culm height and widest leaf width), shows almost all individuals can be determined 
using these two characters alone (Fig. 4). Ranges, means, and standard deviations for this 
larger sample size are presented in Table 4. Mainly depauperate or aberrant individuals 
were not separable with these two characters. These specimens were determined by the 
additional characters as discussed previously with scale color being the most useful. 

Regional differences in the characters were significant only for width and length of 
perigynia (Table 2). Perigynia of C. reznicekii varied by geographical region with slightly 
narrower and shorter perigynia on plants from the northern region compared to the south- 
ern and Missouri/Arkansas regions. These statistically significant differences represent 
only minor variations and have questionable biological significance. 

For the distribution curves for widest leaf width and tallest culm height a total of 


1058 


BRIT.ORG/SIDA 22(2) 


oe. o 
a 
—) 
~ 
ps —) Sd 
is aif Sd 
52 o ° 
Se $ 
joe e @ 
ae ° : 
om af 
mt Qo) + . 
= ‘ a 
—} aa 
=F a o . 
a 2 oe o ¢ = oO 
ca : 
Se + 
oA ? Je 
Be > 
Q 
2 & -0.5 ‘ eo 
KS 0 C. reznicekii ; 
2m * C. nigromarginata s 2° 
< rs a a 
a 
Nn a 
- 
Nn 
-1.5) iE 3 + T 7 T 
-2.5 -1.5 -0.5 ie 1.5 
Axis 1: TaCuHe (-0.961), WiLeWi (-0.747) 
Fic. 3. Non ic multidi ional scaling ordination showi tial relationshir ie ean C. nigromarginata based on 


Apidild J y y 


based On wider WIUCDSL IC dt VIidUe 


. Main characters correlated to each axis 


Taste 2.Ranges, means, and standard deviations for characters ee for the NMDS ordination (Fig 3).Char- 
acter codes are defined in Table 1.Regions listed below are ¢ NE = Virginia/Kentucky north; SE = North 
Carolina/Tennessee south; and AR/MO = Arkansas/Missouri, 
Carex reznicekii Carex nigromarginata 
regions egions 
Characters NE SE AR/MO Allregions NE SE AR/MO All regions 
TaCuLe 2.2-12.2 2.5-13.7 3.7-13.7 2.2-13.7 9.0-27.7 6.6-18.7 7.2-38.0 6.6-38.0 
4+3.1 5.4+3.3 (3235 5:62:33 15.2452 13.3438 175+88 15.3+6.3 

WiLeWi 1.3-2.2 1.3-2.0 1.7-2.2 1.3-2.2 24-40 2.4-3.5 2.5-3.5 24-40 

1.6+£0.3 1.7£03 2.00.2 1.7+0.3 2.9+0.5 2/7404 3.0+0.3 2.9+0.4 
Anle 1.30-1.80 1.30-1.95 1.50-1.75 1.30-1.95 1.50-2.20 1.45-2.25 1.60-2.50 1.45-2.50 

1.50+0.15 1.52+0.17 1600.10 1.5340.15 1.82+0.23 1.84+£0.25 1.98+0.30 1.88+0.26 
StSpExPiSp 0.2-2.5 0.2-2.2 ]-3.7 0.2-3.7 0.7-7.3 1.1-6.0 0.5-5.5 -73 

1.4+0.7 1.2+£0.5 25 [eZ 1.6+0.9 3.4+2.0 2.7+1.6 3.0+1.7 S017 
PeLe 2.75-3.45 2.80-3.70 3.35-3.55 275-3.70 2.70-3.30 2.90-3.60 3.00-3.75  2.70-3.75 

3.16+#0.21 3.23+0.27 3.40+0.10 3.244023 3.1440.18 3.2740.25 3.3740.25 3.264£0.24 
PeWi 0.95-1.10 05-1.20 Henan 0.95-1.20 0.95-1.35 0.90-1.35 1.05-1.25 0.90-1.35 

1.02+0.06 1.13+0.05 1.1340.07 1.09+0.08 1.1140.11 1.14#0.12 1.1240.07 1.12+0.10 
StSpHe 4.60-7.55 4.60-7.85 a ae ie na 5.85-12.10 6.15-11.15 5.35-10.70 5.35-12.10 

6.31+0.89 6.05+1.05 733+ 1.141 841#2.13 7.79+1.58 8.02+1.74 8.07+1.79 
StSpWi eee 1.55 Q.70-1.65 ees ae 1.65 1.05-2.25 1.00-2.00 1.35-1.75  1.00-2.25 

27+0.20 1.20+0.28 1.3140.22 1.2540.23 1.62+0.36 1.504£0.31 1.5240.15 15440.28 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1059 


Tasie 3. Summary of F statistics and probability values from one-way ANOVA between C. reznicekii and C. 
nigromarginata for characters measured for ordination. 


Characters Fi 50 p 
TaCuLe 43.8 <0.001 
WiLeWi 136.2 <0.001 
AnLe 32.6 <0.001 
StSpExPiSp 123 0.001 
PeLe 0.0 0.914 
PeWi 1.6 0.208 
StSpHe 13.3 0.001 
StSpWi 15.7 <0.000 


Taste 4. Ranges, means, and standard deviations for characters measured for the scatter plot graph and charac- 
ter distribution curves (Fig.4,5, and 6). 


C. nigromarginata outside of 


C. reznicekii 


150 specimens 


C. nigromarginata (includes specimens from 
inside and outside the range of C. reznicekii) 
152 specimens 


range of C. reznicekii 
81 specimens 


TaCuLe 1.9=13.7 4.5-51.0 4.5-51.0 
Ss 216 165 2725 19.1+8.0 
WiLeWi 1.2-2.5 1.9-4.5 2.1-4.4 
AZO 2905 3.00.5 


Taste 5.Ranges, means, and standard deviations of particle sizes, pH, and mineral contents of six soil samples. 


Measurements Measurements 

Sand 31-88% Mg = 33.67-221.09 ppm 
57+20 136.10+71.84 

Silt 9-54% Ca 220.28-5509.98 ppm 

#15 1732.66+ 1931.26 

Clay 2-27% Fe 0.64-45.38 ppm 
14+£10 16.00+ 20.97 

pH 4.35-6.51 Al 15.51-317.91 ppm 
549+1.02 111.964117.45 

P 0.89-6.40 ppm Mn 20.43-93.99 ppm 
2.35+2.06 69.17+27.38 

K 24.76-268.13 ppm Zn 0.87-9.21 ppm 
125.79+82.20 3.5443.02 


150 specimens of C. reznicekii and 152 specimens of C. nigromarginata were measured. 
Eighty-one of the C. nigromarginata specimens were from outside of the geographic range 
of C. reznicekii. The distribution curves for widest leaf width and tallest culm height for 
C.nigromarginataand C. reznicekii combined shows a bimodal distribution of these char- 
acters (Fig. 5 and 6). The distribution curves for C. nigromarginata and C. reznicekii as 
separate taxa are overlapping normal or skewed normal curves (Fig. 5 and 6). The distri- 
bution curves for specimens of C. nigromarginata from outside the range of C. reznicekii 
closely mimic the distribution curve for C. nigromarginata throughout its range and also 
clearly contrasts with the distribution curves of C. reznicekii (Fig. 5 and 6). These distri- 


1060 BRIT.ORG/SIDA 22(2) 


60.07 
a 
a0) °C. reznicekii 
*C. nigromarginata 
— 40.0 
2 
y , 
~~ . 
| ee o 
B® 30.0 . oe ° 
: ee . 
= P ter 4, . 
— Sa 
5 20.0 *4, ee Sere .? 
Sd 

~ *$ woe ee 
o #3 % 2 * ¢ 
—| O ry $$+ 33 ¢ aa 
10.0 2 o* $f*, , ° . 

5 s Heteese + 

o = iH ol lUf 

iE 


0.0 T T T rT T T T T T T r T T T rT T —_—_—— 
1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 2.6 2.8 3.0 3.2 3.4 36 3.8 4.0 4.2 44 4.6 4.8 
Widest Leaf Width (mm) 


widest leaf width by t C. nigromarginata and C. reznicekii. 


Cc + lnat f£.4l1 
Fic. 4. Scatter f f g 


80.0 - 
70.0 - : ie : ‘ 
—— C. reznicekii and C. nigromarginata 
-- °C, nigromarginata 
60.0 5 . as 
— --C. reznicekii 
ea — — C. nigromarginata beyond range of C. reznicekii 


Number of individuals 


od vA * 
7 N 
0.0 t T 7 T T LJ v T T T 7 

a 

a a a an Dh a D a an 
7 a 4 . . “0 na e bi + 
YH eo < ~ S ma >: = rs 
= es _ ro] fo) ro] er) oe ‘© a + + 

. ° ye ioe) 
Widest leaf width (mm) 
Fic. 5. Distributi f idest leaf width for C. reznicekii, C. nigromarginata, C. reznicekii and C. ig gi bined, and C. 


f C reznicekii. 


J J r J bal | J 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1061 


100) 
907 
807 C. nigromarginata and C. reznicekii 
2. ~* "°C. nigromarginata 
= 70 
= —""C. reznicekii 
607 ee es ieee 
i C. nigromarginata outside of range of C. reznicekii 
Z 507 
— 
S 407 
Sam 
5 307 
& 
3 207 
Zz. 
107 
“ a 
n — ————" 
() ee a 
SSegRgeege eRRRERRAR RA ZR GB 
4 Foto RA SAREE TS SG 
1 1 
a oO 2 FR aR HH RH GE ESSE 
Tallest culm height (cm) 
Fic. 6. Distributi f ll Imt ig fe reznicekii, C. nigromarginata, C. reznicekii and C. ig gi I 1,and C. 
ig gil I tside the geographic range of C. reznicekii. 


bution curves help to demonstrate that although there is some overlap in the two most 
explanatory characters that separate C. nigromarginata and C. reznicekii, the distinction 
between these two species is not a result of a sampling bias. The two species simply have 
slightly overlapping normal or skewed normal distribution curves. 

Geography 

Carex reznicekii occurs from New England (southern Rhode Island and southern Con- 
necticut) south to southeastern New York, eastern Pennsylvania, South Carolina, and 
Georgia west to southern Missouri and northern Arkansas (Fig. 1). It is quite common in 
the southeastern US. 

The range of C. reznicekii appears to be totally within the range of C. nigromarginata 
and C. umbellata (Crins & Rettig 2002). The distribution of C. nigromarginata extends 
beyond the range of C. reznicekii into southern Ontario, central and western Pennsylva- 
nia, southern Ohio, Indiana, Illinois, central Missouri, central and southern Arkansas, 
Oklahoma, Texas, Louisiana, southern Mississippi, southwestern Alabama, and the pan- 
handle of Florida (specimens examined, Reznicek & Catling 1982; Cusick 1992; Crins & 
Rettig 2002). The distribution of C. umbellata extends well beyond the range of C. 
reznicekii into Greenland, eastern and western Canada, and from Nebraska south to Texas 
(Crins & Rettig 2002). 


Ecology 

Carex reznicekii occupies mesic to dry-mesic mostly deciduous or less commonly de- 
ciduous-pine and/or Ilex opaca Aiton forests. Overstory tree canopy cover often exceeds 
ca. 60% but does not create dense shade. The shrub layer is usually not very dense. The 
herb layer varies from quite diverse to not diverse, and quite dense to not dense. Occa- 


1062 BRIT.ORG/SIDA 22(2) 


sionally C. reznicekii grows in clearings in forests. Even at these sites, it is mostly found 
in the adjacent full-canopy forests. It also occurs along road edges immediately adjacent 
to forests as well as in the forest interior. Overall, it grows predominately in full-canopy 
forests, and unlike some other members of section Acrocystis (e.g., C. nigromarginata and 
C.umbellata), it usually does not do equally as well in more open environments. 

Carex reznicekii often occurs on slopes above drainages from the bases of the slopes 
to mid-slopes and occasionally on the upper slopes or crests. It usually does not occur in 
the actual floodplain of the adjacent drainage unless the drainage is relatively small. 
Additionally, C. reznicekii usually does not occur on upper slopes, especially if erica- 
ceous shrubs and Quercus montana Willd. are dominant. 

Many members of Carex section Acrocystis in eastern North America often grow 
together. Perhaps this is because they grow ina “generalized” habitat (Cusick 1992). Other 
members of section Acrocystis observed growing syntopically with C. reznicekii, often 
within centimeters of it, include C. albicans var. albicans, C. albicans var. emmonsii, C. 
nigromarginata, C. pensylvanica Lamarck, C. tonsa var. tonsa, C. tonsa var. rugosperma, 
and C. umbellata. Carex nigromarginata occurred as the most frequent associate, being 
found with C. reznicekii at 19 out of the 31 sites visited. Despite C. nigromarginata fre- 
quently growing syntopically with C. reznicekii, some habitat differences between the 
two were observed. Carex reznicekii usually grew without or with low densities of C. 
nigromarginata in more calcareous or richer sites as well as lower on slopes. Carex 
reznicekii usually did not occur or occurred in lower densities with C. nigromarginata at 
more acidic sites, higher up on slopes, or in more open disturbed habitats. 

Soils are deep or less frequently shallow over bedrock. Bedrock present at sites in- 
cludes diabase, limestone, shale, and sandstone. Soil samples showed a wide variety of 
soil textures ranging from sandy to sandy loam to silty loam to sandy clay loam to clay 
loam. Table 5 summarizes the ranges, means, and standard deviations of values for par- 
ticle size, pH, and mineral content of the six soil samples. 


IDENTIFICATION KEY FOR MEMBERS OF CAREX SECTION ACROCYSTIS 
EAST OF THE ROCKY MOUNTAINS OF NORTH AMERICA 


This key works best with ample specimens in mature fruit. Depauperate or especially 
robust specimens may not key correctly. Rhizomes help in identification, but collectors 
often fail to collect them. The key attempts, where possible, to get around relying on rhi- 
zome characters. Parts of the key dealing with C. communis varieties and C. inops ssp. 
i aii are adapted from Crins and — _ 


Ir fare fea ‘ lee 


. Cul pedt near base lige eae 
times s difficult ie assess pecsuee ne ere can hidden . sheathes of bracts and leaves 
until they emerge adjacent to distal half of culms); pistillate spikes arising from tee half of 
culms mostly 0- 
2. Pistillate scales om slightly shorter to longer than the perigynia; terminal staminate spikes 
with or without approximate pistillate spikes; bracts at base of proximal-most non-basal 
ou spikes (if present) mostly shorter than to sometimes slightly longer i. inflores- 
ences; bases of old leaves often very fibrous 
: Perigynia (2.2—)2.3-3.2(-3.3) mm long, beaks a )0.5-1.0 mm long Carex umbellata 
3. Perigynia (3.0-)3.1-4.7 mm long, beaks 0.9-2.1 mm long. 
4. Perigynia often mostly glabrous; young — short, thick, often smooth adaxially, and 
ri Carex tonsa var. tonsa 
4. Perigynia usually pubescent; young leaves long, thin, scabrous adaxially, and not rigid 
rex tonsa var. rugosperma 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1063 


2. Many pistillate scales shorter than mature perigynia; terminal staminate spikes almost al- 
wae ae at ce one sabe inane Piauliete spike; Piags a nase of shee most 


y longer s notor only slightly 
fibrou 
> ne (2.2—-)2.6-3.1(—3.2) mm long, beaks 0.4-0.8 mm long, beak teeth up to 0.2 mm 
long; pistillate scales often shorter than the body of mature perigynia; staminate spike 
3.5-5.9(-6.7) mm long; rhizomes slender; plants loosely caespitose Carex deflexa 
Hornem. var. deflexa 
5, Perigynia 3.1-4.2 mm long, beaks 0.9-1.7 mm long, beak teeth 0.2-0.5 mm long; ae aia 
scales ond longer than the body of mature perigynia; staminate spikes 4.2-11.6 m 


lon j zomes ice) ut; plants densely Ca 


ng;r spitose pss rossii Boott 
. Culms with ectie pies: short- seauneulaie to sessile (proximal-most non-basal spikes oc- 


casionally elongate-pedunculate) and arising from distal half of culms [occasional culms will 
have one pistillate spike with an elongate peduncle arising from base of culms ( 
represent hybrids)]; pistillate spikes arising from distal half of culms mostly 2-4 
6. Longest staminate spike at least 12.3 mm long (use ample specimens); rhizomes elongated 
(except C. albicans var. albicans, C. communis, and C. novae-angliae: these ane out in either 
lead of 6): perigynia bodies globose to sree as long as or longer than 
7. Widest ae equal to or less than 1.2 mm wide; widest leaf ee 
wide; perigynia bodies ellipsoid, longer than wide. 
8. Widest ae anes 1.8 mm wide; proximal internode of inflorescence 5.2—29(—40) 
m long; staminate ae peduncles 1.2-8.0(-8.6) mm long; proximal-most pistillate 
ae peduncles 0.8-9.8 mm long; spike above the proximal-most spike subtended by 
twh ftenh longat pistillat | tred, i with s 


these may 


1.1-2.7 mm 


me 


ed-brown Carex novae-angliae Schwein. 
8. Widest leaf blades 1.3-2.7 mm wide; proximal internode of inflorescence 1.0-13.2( 
0.1) mm long; staminate spike peduncles 0.3-4.5(-8.8) mm long; proximal-most pis- 
tillate spike peduncles 0.3-2.2(-3.3) mm long; spike above the proximal-most spike 
subtended 2 a bract which usually has at most a short awn; pistillate scales some- 
times partly r 
9. Rhizomes Seeaes and spreading 
9. Rhizomes short and ascending to erect 
iS Widest poy ae cag anes 2mm wide; wi widest feat blades 1.5-6.1 
out 


Carex albicans var. australis (Bailey) Rettig 

x albicans var. albicans 
mm wide; 

wide to slightly longer 


= 


than W 
10. ene leaves (2.3-)3.0-6.1 mm wide; plants with large caespitose clumps; not colo- 
nial; rhizomes short and ascending to erect; bracts subtending the spike above the 
proximal-most with an elongate awn tip that arises from scarious margins that are 
truncate to convex at summit; staminate spikes 0.9-2.2(-2.5 wide 
1. Perigynia beak teeth 0.1-0.2(—0.3) mm long; pistillate scales usually 1.6 mm wide 
or less; widespread 
11. Perigynia beak teeth usually more than 0.2 mm long; pistillate scales usually more 
than 1.6 mm wide; Georgia and South Carolina 


Caren communis var. communis 


Carex communis 
var. amplisquama i Rettig 
10. Widest leaves less than 3.0(-3.5) mm wide; plants with small caespitose clumps; colo- 

nial; rhizomes spreading and long; bracts subtending the spike above the proximal- 
most with a short or no awn tip that arises from lee margins that mostly taper 
into the distal part of the bract; staminate spikes (1.1-)1.5-3.2(—5.2) mm wide. 
12. Perigynia 1.1-1.6(-1.8) mm wide. 

13. Perigynia beaks 0.3-0.8(-0.9) mm long, beak/body ratio 0.13-0.50(-0.56) 


Carex pensylvanica 


13. Perigynia beaks 0.9-1.6 mm long, beak/body ratio 0.50-0.92 
Widest leaf blades greater than (1.7-)2.0 mm wide; northeastern North 


Am Carex lucorum Willd. ex Link var. lucorum 
14. Woes leaf blades less than 2.0 mm wide; southern Appalachian 
Mountains 


Carex lucorum var. austrolucorum Rettig 


1064 


6. 


BRIT.ORG/SIDA 22(2) 

12. Perigynia (1.6-)1.7-2.2 mm wide Carex inops ssp. heliophila 
Longest staminate spike less than 12.3 mm long (use ample specimens); rhizomes not elon- 
gated (except C. floridana and C. albicans var. australis: the latter keys out in either lead of 6) 


perigynia bodies ellipsoid, longer than wide (except C. deflexa and C. communis: the latter 
keys out in either lead o 


15. Pistillate scales shorter than body of mature perigynia they subtend. 
16. P 


roximal internode of inflorescence 5.2-29(-40) mm long; staminate spikes (4.6— 
5.1-12.1 mm long 


Carex novae-angliae 
16. Proximal internode of inflorescence 1.3-8.5(—14.5) mm long; staminate spikes 3.5 
8.3(-9.2) mm lon 
7. Culms mostly surpassed by leaves; perigynia (2.2-)2.6-3.1(-3.2) mm long Carex 
deflexa var. deflexa 


Culms surpassing leaves; perigynia (3.0-)3.1-4.2 mm long Carex peckii Howe 


15. ee oan longer than body of mature perigynia they subtend. 
18. Widest leaf blades (2.3-)3.0-6.1 mm wide; proximal internode of the inflorescence 


(5.9-)7.5-36.0 mm long; bract subtending the spike above the proximal-most w 
an oe awn tip that arises from s 
| j ia hod) re 


arious margins that are truncate to convex at 
tol lly ellipsoid g to slightly longer 


to broadly ely about 


than wide. 


9. Perigynia beak teeth 0.1—0.2(—0.3) mm long; pistillate scales usually 1.6 mm wide 
or less; widespread 


communis var. communis 
9. Perigynia beak teeth usually more than 0.2 mm long; pistillate scales usually more 
t 1.6 mm wide; Georgia and South Carolina arex communis 


var.amplisquama 
18. Widest leaf blades 1.1-4.5 mm wide (if widest leaves are over 2.7 mm wide then 


proximal internode of the inflorescence 0.8—5.4 (8.0) mm long); bract subtending 
the spike above the proximal-most with or without an elongate awn tip that arises 
from scarious margins that mostly taper into the distal part of the bract (occasion- 
ally truncate to convex at summit); perigynia bodies ellipsoid, longer than wide 
20. oe leaf blades 1.1-1.8 mm wide; proximal internode of inflorescence 5.2— 
-40) mm long; staminate ae oooada at 
most pistillate spike peduncles 0. the proximal-most 
spike subtended by a bract waied often has an elongate awn; pistillate scales 
without true red, sometimes with some red-brown Car 

. Widest leaf blades 1.2-4.5 mm wide; proximal eae of inflorescence 0.8- 
13.2(-20.1) mm long; staminate spike peduncles 0.1—4.5(-8.8) mm long; proxi- 
mal-most pistillate spike peduncles 0.0-2.2(—3.3) mm long;spike above the proxi- 


mal-most spike subtended by a bract which usually has at most a short awn 
pistillate scales sometimes with s 


1.2- ool 8.6) mm long; ‘Proximal: 


NO 
(Se) 


arex novae-angliae 


21. Longest leaf blades up to 2.2(- oe cme longas th 


median culm length; 
see 2.2—3.2(-3.3) mM long; proximal internode of inflorescence 1.0- 
13.2(-20.1) mm long; staminate spike peduncles 0.3-4.5(-8.8) mm lo 
22. Rhizomes elongate and spreading 
22. Rhizomes short and ascending to erect 
23. Staminate spikes (5.5-)6.6-1 


Carex hes var. australis 


2.2(-14.5) mm long; middle and upper 
staminate scales mostly with midrib faint, not raised, not scabrous 
towards tip of scale, and mostly without a minute awn arising just 
below tip of scale ex albicans var. albicans 
. Staminate spikes 4.0-8.7(—9.3) mm long; middle and er stami- 
nate scales mostly with midrib more prominent, raised, antrorsely 
scabrous towards ae of scale, and with a minute awn ary from 
just below tip of sc C 


NO 
WW 


No 
mart 


var. emmonsii 
ieee Hee ee )2.5 times as long as the median culm length; 


perigynia (2.5—)2.7-4.0 mm long; proximal internode of inflorescence 0.8 
5.4(-8.0) mm long; staminate spike peduncles 0.1—0.8(—1.6) mm lon 
24. Rhizomes horizontally spreading to ascending and elongate; mostly at 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1065 


least some stigmas 2; mostly at least some achenes with only 2 pale 
nerves; sheathes of old leaves at base of plant usually weakly ote 
Carex floridana 
24. Rhizomes ascending to erect and short; stigmas 3; achenes with 3 pale 
nerves; sheathes of old leaves at base of plant often strongly fibrous. 
25. Widest leaf blades 1.2-2.2(-2.5) mm wide; culms often about equal 
length, the longest 1.9-9.9(-13.7) cm long; any reddish color on pis- 
tillate scales below the distal tip not extending laterally Helin near 
the margin to the green or brown longitudinal mid-strip 
side of mid-vein; the longest staminate spike up to 3.7 mm oiler 
oS the distal most lateral cass am staminate spikes 3.3-8.3( 
m long X 0.4-1.6(-1.8) m e Carex reznicekii 
: fe leaf blades (1.9 cee mm sere culms often of quite vari- 
able length, the longest (4.5-)6.6-38.0(-51.0) cm long;at least some 
pistillate scales often with reddish to purplish/black color below the 
distal tip extending laterally from near the margin to the green or 
brown longitudinal mid-stripe on either side of lea long- 


N 
[oa] 


est staminate spike (0.0-)0.5-5.1(-7.3) mm taller t! stal most 
lateral spike apex; staminate spikes 4.2-10.9(-12.0) mm x 0 
2.4(-2.8) mm wide Carex Taremerenat 


Representative specimens of Carex reznicekii: poemce sabes with a single artes were measured for the 
ordination (Fig, °) ale ones marked witha single or d 


tk 1€ sCallel plot graph and 


character ibution curv Fig 4 5a nd 

.A, ALABAMA. Dekalb Co.: near top of Red Mountain ae NW of Fort Payne, 21 Apr 1943, Harper 3918 
(MO, ust ), Harper 3919 (BH**, GH, US). Houston Co.: vicinity of Columbia, Thomas Creek up to bluff along 
Ommussee Creek, 12 Mar 1998, MacDonald 10970 (BRIT**). Jackson Co.: ca. 3 mi W of Carns, along E side of 
county Route 33, 22 Apr 2002, Naczi 9034 (DOV™*); just E of Route 35, 0.1 mi S of junction with Route 40, 21 Apr 
2004, Werier 2075 (BH*), county Route 33, 8.05 mi by road SE of junction with Route 79, 21 Apr 2004, Werier 
2081 (BH**, MICH, NCU). Limestone Co.: by 1-65, ca. 5 mi N of US 31 turnoff to Athens, 26 Mar 1986, Kral 73127 
(BRIT **). Morgan Co.: by US 31,1 mi S of Lacon, 12 Apr 1970, Kral 38342 (BRIT*). tascalovss Co.: by AL 69,9 mi 
N of Moundville, 1 Apr 1984, Kral 71138 (BRIT**). ARKANSAS. Baxter Co.: Cole F | pple Turkey Road, 
26 Apr 1993, Hyatt 5243 (MICH*). Independence Co.: 6.95 mi S of Sharp Co. Line (in Cave ae on US Hwy. 167,9 
Apr 1994, Hyatt 5838 (MICH*), Hyatt 5839 (MICH*). Izard Co.: ca. 0.5 mi SE of Baxter Co. on AR Hwy. 5, 18 Mar 
1998, Hyatt 7879 (MICH**). Johnson Co.: [Ozark National Hore st], Horsehead Lake Recreation Area, western most 


picnic area, 12 Apr 1996, Hyatt 6896 (MICH*). Marion Co.: Buffalo N | River, 100-200 m NE of Buffalo River 
Bridge of state Hwy. 14, 10 Apr 1994, Hyatt 5844 ne Newton Co.: ‘= of Harrison, 0.8 mi N of Harp Creek, 
along Hwy. 7,14 Apr 1984, Castaner 7992 (DOV**). Stone Co.: Mountain View vicinity, about 10 mi SSE on AR 


Hwy. 5,8.5 mi N of Cleburne Co. line, 18 Mar 1998, Hyatt ae a), CONNECTICUT. New Haven Co.: ee a 
New Haven, West Rock Ridge, a little N of “Judges Cave,” 20 May 2005, Werier 2533 & Haines (BH** 
Kent Co.: above Choptank River floodplain at Sandtown landfill, N of Sandtown, 30 Apr 2003, pee = 
(DOV*). [New Castle Co.]: Mount Cuba, | May 1898, Bicknell 3255 (NY*); near Centreville, 28 Apr 1876, Commons 
sn. (PH**). Sussex Co.: 3 mi E of Milford, along S side of Beaverdam Branch, ca. 0.2 mi W of its crossing by road 
200, 21 Apr 2003, Naczi 9521 (BRIT, DOV™). [Sussex Co.]: Laurel, 25 Apr 1882, Smith 717 (US*™). [DISTRICT OF 
COLUMBIA]. [Washington Co.]: Washington, 25 Apr 1897, Steele s.n.(GH**); District of Columbia and vicinity, 7 
ia 1899, Williams s.n.(NY**), Anacostia to Twining City, 7 Apr 1918, Leonard 139a (US**). GEORGIA. Clarke Co.: 
of Athens, ce of Rock and Shoals Road off Barnett Shoals Road, 24 Feb 1981, Manhart 202 (BRIT**, MICH**, 
a NYS,). Clay Co.: 1 mi E of Fort Gaines, 27 Mar 1949, Thorne G& Muenscher 9312 (GH**). Dade Co.: Sitton’s 
Gulch, Lookout Mountain, 2 mi SE, Trenton, 17 Mar 1939, Hermann 10192 (GH*, NY*™). Hall Co.: Chipowee Woods 
(Elachee Preserve), 19 Mar 2005, Hyatt 11802 (BH**), 9 Apr 2005, Hyatt 11807 (BH**). Randolph Co.: near Griers 
Cave, N of Cuthbert, 3 Apr 1948, Thorne & Muenscher 7910 (BH**). Walker Co.: W side of GA Route 136, 1.6 mi by 
road W of GA Route 193, 0.4 mi W of Cooper Heights Community, 8 Apr 2000, Hill 32220 ot Ww a 
136, 1.6 mi by road W of junction with Route 193, 20 Apr 2004, Werier 2069 (BH*, MICH, NCU). KE 
Butler Co.: Roundabout Swamp, at end of Biggerstaff Road, off Hwy. 1153, 22 Apr 1993, ees 5360 es 
Edmonson Co.: Mammoth Cave National Park, Ugly Creek S eae, with Little Jordon Cemetery Road, 23 
Apr 1991, rane aa 4622 (BRIT**). Rowan Co.: Lic ek ork, 4 1938, Braun 1758 (US**). MARYLAND. 
Caroline Co.: Tuckahoe State Park, off Cherry Lane l mi qorniae with Mason Branch Road, 


J wl 


1066 BRIT.ORG/SIDA 22(2) 


14 Apr 1997, Frye 779 (MICH**), 2 May 2004, Werier 2176 (BH*), Werier 2178 (BH**, MICH, US); Tuckahoe State 
Park, entrance to C Lane picnic area., 2 eo reall Werier 218] (BH**, MICH, US). Cecil Co.: along Bohemia 
Creek, Middle Neck, 7 May 1932, Long 37288 (PH**). Dorchester Co.: N side of Puckum Branch, W of Puckum 
Road, N of Eldorado, 15 oe 2003, McAvoy 5797, ae GE Biot (DOV*). Kent Co.: above the “Big Marsh,” W of 
Betterton, 27 se 2003, McAvoy 5739 (DOV**). Montgomery Co.: Plummer’s Island in the Potomac River, n 
Cabin John, 10 May 1915, Van Eseltine 325 (BH**). Wicomico Co.: S side of Bear Swamp Road, SW of See ‘ 
Apr 1999, a 4268 (DOV**). MISSISSIPPI. Lafayette Co.: NE of Oxford, 5.4 mi NE of junction of MS 7 and 30 
on MS 30 then ca. | mi S on Hopewell Road, 12 Apr 1997, Bryson 15732 (BRIT, DOV**). Lowndes Co.: 2.5 mi N of 
Steens, N side of Vaughn Robertson Road, 20 Apr. 1986, Bryson 4217 & Curtis (MICH, MO), 12 Apr 1987, Bryson 
5341 (MICH*™*). MISSOURL. Shannon Co.: above intermittent tributary of E branch of Dennis Hollow, 13 May 
1997, Brant 3696 (MO**): Rocky Creek Falls, 24 Apr 1983, Merriott 61 (MICH*). NEW JERSEY. Burlington Co.: 
Bordentown, along Crosswicks Creek, 27 Apr 1910, oe 3103 (NY**). [Gloucester Co.]: Swedesboro, 25 Apr 1917 
Mackenzie 7559 (NY**). Sussex Co.: Andover Township, SW of Lake Mohawk, just N of Route 613, 0.5 mi by road 
W of junction with Route 617, 4 May 2004, Werier 2191 (BH*™, iene NY); Stokes State Park, S of county Route 
636, 0.25 mi E of US Route 206, 4 May 2004, Werier 2194 (BH*, MICH, NY). [Warren Co.]: between Pike and 
Pequest River ca. 1/4 to 3/8 mi E by slightly NE of Buttsville, 18 May 1919, Pretz 9602 (PH**). NEW YORK. New 
York Co.: Manhattan Island, Inwood, 27 May 1893, Bicknell 1237 (NY**). Orange Co.: Town of aa SE end of 
sor es aan N of the village of Greenwood Lake, SW of Route 17A, 7 May 2003, Werier 1507 & Barbour 
(BH*). Ulster Co.: Town of Saugerties, 1.0 km E of Old Dutch Road, 1.1 mi S$ eg road from Route 32, 22 May 2006, 
Werier se Barbour & Barbour (BH). NORTH CAROLINA, Davidson Co.: 1 mi E of Yadkin River near NC 4 
Apr 1956, Radford 9476 (NCU*™*); S of Route 49, 0.6 mi E of bridge over Yadkin River, 14 vy 2004, Werier 1995 
(BH*, MICH), Werier 2000 (BH**, MICH, NCU), Werier 2001 (BH**, MICH). Macon Co.: Nantahala National For- 
est,ca.5 mi E of Franklin, then 5 mi N of Ellijay Road, 16 Apr 2000, Hyatt, ae a Me Millan 9112 (MICH**); W of 
Ellijay Road and Ellijay Creek, 3.7 mi by road NE from Route 64, 19 Apr 2004, Werier 2048 (BH*, MICH, NCU); E 
of Route 2, 5.4 mi by road NE from Route 64, 19 Apr 2004, Werier 2052 (BH**, DOV, MICH). Madison Co.: 0.5 mi 
WNW of Marshall on NE side of ave n Broad River, 26 Apr 1958, Ahles 38966 & Duke (NCU**). Mecklenberg 
Co.: 2.4 mi SE of Davidson on NC 73, 22 Apr 1958, Ahles 38691 & Britt (NC — Co.: Phills Creek, N side, 
3 mi W of Chapel Hill, 18 Apr 1940, Radford & Stewart 469a (NCU**). Sw : Nantahala National Forest, 
Wessen vicinity, S side of US Hwy. 19, 10 Apr 2000, Hyatt 9071 (MICH**). PENNSYLVANIA. Berks Co.: 5/8 mi ENE 
of Walnut & 14th Street, Reading, 20 May 1943, Berkheimer 3563 (NYS*). [Lancaster Co.]: Lancaster, aaa [no 
collector] s.n. (MICH**); [no location information], [no date], [Muhlenberg ?]s.n. (PH**), [no location information], 
[no datel, Mu ae: s.n.(PH**). RHODE ISLAND. [Washington Co.]: South Kingston, 13 May 187 pee ie 1776 
H CAROLINA. Chester Co.: near Catawba River at Great Falls, 15 Apr 1961, Radford 43511 (NCU™). 
ame Co.: W of Catarrh, just E of Lynches River, and just S of Route 903, 17 Apr 2004, Werier 2013 eee 
MICH). Fairfield Co.: 2 mi NE of Altston, 15 Apr 1961, Radford 43473 (N NCU**). Greenville Co.: E of Marietta, small 
road, 20 m SE from Route 414, 2.1 mi by road NE from Route 276, 18 Apr aoa - erier 2029 (BH**, MICH). Horry 
Co.: 2 mi S of Myrtle Beach, 18 Apr 1932, Weatherby & Griscom 16461 (PH **) Lancaster Co.: near Catawba 
River, 2 mi NE of Great Falls, 15 Apr 1961, Radford 43524 (NCU): E of Great 7. anda little bit E of the ees ba 
River, 17 Apr 2004, Werier 2023 (BH**, MICH), Werier 2027 (BH**); E of US 601, S side of Flat Creek, 21 Apr 19 
Williamson F138 & Ahles (NCU*™), 16 Apr 2004, Werier 2010 (BH*, MICH). Pickens Co.: SC Department of ee 
ral Res stern tract of Lake Jocassee, S of Mill Creek entrance, 19 May 1998, Waldrop W-274 & Townsend 
(MICI > just E of tributary to Oolenoy River and SSR 69, 0.1 mi by road N from Route 288, 2.5 mi by road E from 
Route 178, 18 Apr 2004, Werier 2038 (BH*™*, MICH), Werier 2041 (BH**, DOV NCU). TENNESSEE. Franklin Co.: 
x 6 road mi S of Huntland, along E of oo 97, 2.6 road mi S of junction of Routes 97 and 122, 24 tel 1998, 
zi 7249 & Ford (VPI*™). Grundy W of state Route 50 (Elk Head Rd.), 5.7 mi by road SW from junc- 
tion an Route 108, 22 Apr 2004, ara oe (BH*, MICH); E of Route 56 at hair pin turn, 6.6 mi by ee S from 
Warren Co, line, 22 Apr 2004, Werier 2098 (BH™, ea Co.: W side of the Cumberland River at its 
junction with Cub Creek, 19 Apr 1998, Phillippe 29579 (MICH**); NE of Cub Creek just before junction with 
Cumberland River, 23 Apr 2004, Werier 2102 (BH**, MICH). Jefferson Co.}: French Broad River, near eee aa 
1844, Rugel s.n. oe NY**). [Knox Co.]: Knoxville, Cherokee Bluffs, 9 Apr 1930, Underwood 5 (GH*™*). Marion 
Co.: Send of Little Cedar Mountain, W side of Nickajack Lake, and 2.8 km N of Interstate 24 bridge W i Haletown, 
8 Apr 2000, wea 10418 (MICH**, VPD; E slope of Little Cedar Mountain, ca. 3.5 km N of interstate 24 bridge 
over Nickajack Lake, NNW of Haletown, 8 Apr 2000, Wieboldt 10423 (MICH™*, VPI). Maury Co.: bluffs of Duck 
River ca. 2.5 mi SE of Columbia and 1.2 mi E on Sowell Pike from TN 50, 13 Apr 1974, Kral 42463 (BRIT**, MO). 
Polk Co.: N of Ocoee Lake and Route 64, 1.7 mi W of junction with Route 314, 20 Apr 2004, Werier 2062 (BH*), 20 
Apr 2004, Werier 2063 (BH**, MICH). Putnam Co.: bluffs along Caney Fork River by I-40, 10 Apr 1992, Kral 80112 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1067 


& Moore (BRIT**). Rutherford Co.: just N of Luvergne, 9 Apr 1976, Kral 57547 (BRIT**, MO). Sevier Co.: N of Little 
Cove a Road, 4.8 mi by road E of Wear Valley Road, 27 Apr 2004, Werier 2132 (BH**, MICH). VIRGINIA. 
Accom .: Sside of road to Sinnickson, 18 Apr 1999, McAvoy 4266 (DOV**). elbow arle Co.: Mount Alto, near 
nae 21 Apr 1973, Stevens 6523 (NCU*), 21 Apr 1973, Stevens 6525 (NCU*™*). [Albemarle Co.]: Monticello 
Mountain, the Grove, 21 Apr 1980, Wieboldt M-26 (PH**). Alleghany Co.: N of Route 18, N of small creek which is 
a tributary of Potts Creek, 0.6 mi by road S from Route 657, 30 Apr 2004, Werier 2153 (BH**, DOV, MICH, VPD, 30 
Apr 2004, Werier 2155 (BH**); 30 Apr 2004, Werier 2156 (BH**, MICH, VPD. Botetourt Co.: NW of Route 615, 0.7 
mi by road N of junction with Craig Co. line, 29 Apr 2004, Werier 2143 (BH**, MICH, MO, NCU, VPD, 29 Apr 2004, 
Werier 2146 (BH**). Brunswick Co.: along Hwy. 670, 2.3 mi NE of its junction with Hwy. 46, 2 May 1986, Rettig 
1523 (NYS**); Route 670, 2.2 to 2.3 mi by road N of Route 46, 11 Apr 2004, Werier 1983 (BH**, MICH, VPI); along 
Hwy. 712, 7.6 mi N of its junction with Hwy. 58 in Edgerton, 2 May 1986, Rettig 1525 (MO, NYS*™*); 7.7 to 7.8 mi by 
road N of Route 712 from Route 58 in Edgerton, 1] Apr 2004, Werier 1972 (BH**, MICH, VPD, Werier 1979 (BH**). 
Buckingham Co.: <— oe of David Creek, ca. 1 mi NE of Bent Creek, 9 May 1983, Wieboldt 4635 & Britten 
(MO**, VPI. C e Con rth Anna River, 10 Apr 2004, Werier 1954 (BH**), Werier 1958 (BH**), Werier 
1960 (BH*™*, en ‘Werier 1961 oe MICH). Chesterfield Co.: 7 mi WNW of Winterpock, along S side of Route 
360, 0.2 mi E of Appomattox River, 30 Apr 2005, Naczi 10792 (BH, DOV**, MICH). Culpepper Co.: 2nd peak, Buz- 
zard Mountain, 11 May 1941, Allard 8589 (VPI*). Fauquier Co.: C.F Phelps Wildlife se ie Area, ca. 1.9 mi 
SW ss Kelly Ford, E of Rappahannock River, | May 2004, Werier 2168 (BH**, MICH, NCU, VPI). Henry Co.: 
along Smith River about 0.9 mi below state Route 636 bridge and ca.5 mi NNE of Ridgeway, 28 Apr 1987, lee 
see Wiebol idt oe Isle of Wright Co.: near Walters, 6 Apr 1938, Fernald & Long 7775 (GH, PH**). King 
e Co.: 2 mi SSW of Owens, along E side of Route 301, 1.5 mi SW of its junction with Route 218, 29 hs 2005, 

ae 10778 ca ee Montgomery Co.: N of state Route 637 at NW end of Coffee Valley, ca. 7 km E of Ellett, 
1l May 1993, Wieboldt 8509, Ludwig, Rawinski, et al. (DOV**, MICH*™, VPD; N of Coffee Hollow at NW end of 
hollow, NE of Seneca Hollow Road, 28 Apr 2004, Werier 2138 & Wield boldt (BH**, MICH). New Kent Co.: 4m 
of Seu along E side of Route 33, 0.5 mi SW of its junction with Route 273, 1 May 2005, Naczi 10822 on 
DOV**, MICH). Newport News City: behind baseball field on Monroe, Fort Eustis, 22 Apr 1974, Appler 1064 & 
ee) (NCU). Patrick Co.: along Spoon Creek ca. 10 mi SE of Stuart, 27 Apr 1983, Wieboldt 4619 (MO**, VPD. 

Pittsylvania Co.: along N-facing bluff of the Staunton River ca. 4 mi N of Straightstone, 23 Apr 1985, Me 
5472 (MICH*™, VPI} aoe Mountain, ca. ; km NE of Leaksville Junction on Southern Railroad, 28 Apr 1 
Wieboldt 8454 (DOV™, MICH**, VPD. P . Powhatan hee Wildlife Management Area, NW of i con, 
11 Apr 2004, Werier 1964(BH* ‘ -MICHD, ele eon MICH, VPD. Prince Edward Co.: 7 mi SSW of Farmville, 
along tributary of Briery Creek, ca.0.3 mi N of dam olf Briery eet. Lake, S of road 790, 30 Apr 2005, Naczi 10811A 
(DOV**). Prince George Co.: Coggins Point, 4 Apr 1939, Fernald & Long 9699 (GH, PH**). Shenandoah Co.: S end 
of Short Mountain, Massanuttens, ca. 3 mi SE of Mount Jackson, | May 1982, Wieboldt 4284 & Davenport (VPI*™). 
Southampton Co.: by the Nottoway River near Davis School, Indian Road, NW of Courtland, 4 Apr 1940, Fernald, 
Long, & Pease 11654 (GH*™, PH**). Surry Co.: W of Claremont, 19 Apr 1942, Fernald, Long, & Abbe 14117 (GH, 
PH**), ca. 1/2 mi NE of reactors at VEPCO Surry Nuclear Power Station, 13 Apr 1974, Ware 5391 & Wieboldt (NCU). 
Sussex Co.: 4 mi S of Stony Creek, 5 Apr 1937, Fernald & Long 6952 (GH, PH**). Wythe Co.: Rock Creek 2.6 km E 
of community of Cripple Creek, 6 May 2004, Wieboldt 11390 (VPI*™*) 


oe 


APPENDIX A 


Specimens of Carex nigromarginata measured for the ordination (Fig. 3) are marked with 
a single asterisk; the ones marked with a single or double asterisk were measured for the 
scatter plot graph and character distribution curves (Fig. 4, 5, an 


CANADA. ONTARIO. Norfolk Co.: Long Point, Squires Ridge, 10 1/2 to 13 mi from base of point, 25 May 1980, 
ae icek 5496 (MICH**); Long Point, middle of a Creek Ridges to mee Bay, 15 to 19 mi from base of point, 
5 Jun 1980, Reznicek 5636 (MICH**). U.S.A. ALABAMA. Autauga Co.: Autauga County 9, W of Ivy Creek, ca.0.5 
mi E are with Autauga County 15, 16 ee 1980, Wiersema 1537 (BRIT**). Butler Co.: by county ei ae ol 
Pigeon Creek, 11 Mar 1991, Kral 78407 (BRIT**); by I-65,4 mi N of exit to AL 106, 26 Mar 2002, 14 (BRIT 
Dallas Co.: ca. 1 mi WNW of Cahaba, 31 Mar 1972, Kral 45340 (BRIT**). Escambia Co.: N side a Rolle 55,ca.05 
mi W of Route 41,19 Mar [19]82, Wilhelm 10013 (BRIT**). Jackson Co.: just E of Route 35,0.1 mi S of junction with 
Route 40,21 Apr 2004, Werier 2072 (BH”*); off of county Route 33,8.05 mi by road SE of junction with Route 79, 21 
Apr 2004, Werier 2079 (BH**). Monroe Co.: E bank of Alabama River above landing at Haines Island, 15 Apr 1980, 
Wiersema 1482 (BRIT**). Morgan Co.: by US 31,1 mi.S of Lacon, 12 Apr 1970, Kral 38342 (BRIT**). Perry Co.: Hwy. 
219,3.0 mi S of Bibb County line, 9 Apr 1986, Rettig 1443 (BRIT**). ARKANSAS. Bradley Co.: Warren Prairie Natu- 


1068 BRIT.ORG/SIDA 22(2) 


ral Area, SE of Warren, 17 Apr 1986, Rettig 1494 (MO**); Warren Prairie, ca. 15 mi W of Monticello, 3 Apr 1983, 
arc 2962 (BRIT*™); about 1 mi W of pe 18 Mar 1995, ean 6161 (MICH**). Calhoun Co.: Hwy. 160 
mi W of Bradley County, 18 Mar 1995, Hyatt 6166 (MICH**). Clark Co.: Caddo Valley vicinity, 1 May 1995, 
yt oe (MICH**). Columbia Co.: US Hwy. 79 at Ouachita ees Te 18 Mar 1995, Hyatt 6173 (MICH**), 
Crawford Co.: just S of Rudy, 20 Apr 1985, ae 8218 (MO*); Road 220 just S of Lee’s Creek, 20 Apr 1985, 
Castaner 8224 (MO**). Drew Co.: Monticello, A.& M. College Farm, 21 Apr 1937, Demaree 14558 (MICH**); 2.8 mi 
S of Lincoln County on US Hwy.425,29 Mar 1997, Hyatt 7287 (MICH**). [Garland] Co.: Hot Springs National Park, 
Mar 1930, Connell s.n. (MO*). Howard Co.: Saline River at its junction with Hwy. 84, 15 Apr 1986, Rettig 1474 
(MO**). Miller Co.: along Hwy. 237,0.7 mi N of its junction with Hwy. 160, 14 Apr 1986, Rettig 1470 (BH**); Hwy. 
237,0.3 mi S of junction with Hwy. 134, 14 Apr 1986, Rettig 1472 (BRIT**). Nevada Co.: AR Hwy. 4 at Ouachita 
County line, 18 Mar 1995, Hyatt 6172 (MICH**):; by I-30, 3.3 mi SW of junction with AR 26, 16 Apr 1999, Kral 88225 
BRIT**). Quachita Co.: NW of Camden, AR Hwy.4 loop, 1.6 mi E of AR Hwy. 376 junction, 18 Mar 1995, Hyatt 6169 
(MICH**). Pike Co.: 0.3 mi S of Montgomery County on AR Hwy. 369, 15 Apr 1996, Hyatt 6938 (MICH**). Polk Co.: 
Ouachita National Forest, Along Forest Service Road 38 to Shady Lake, 3.2 mi N of its junction with Hwy. 246, 15 
Apr 1986, Rettig 1477 (MO**); Ouachita National Forest, along Forest Service a 38, 1.0 mi N of entrance to 
Shady Lake at j EOL with pores Service Road 508, 15 Apr 1986, Rettig 1480 (MO**). Pope Co.: Long Pool 
Recreation Area,O rest, above Big Piney Creek, 16 Apr 1986 cee 1491 (MO*). Sevier Co.: Saratoga 
So 5 Apr Reese (MICH*™); DeQueen, 1.9 mi N on US Hwy. 71,23 Apr 1995, Hyatt 6949 (MICH). 
ott Co.: Ouachita National Forest, approximately 5.5 mi S, 1.5 mi E of junction of Hwy. 80 and business loop 
pee 71 in Waldron, 9 Apr 1999, Morse 2843 (BRIT**); Ouachita National Forest, approximately 2.5 mi S, 3.0 miW 
of Boles, 10 Apr 1999, Morse 2904 (BRIT**). Sharp Co.: N of Cave City on 167,2 mi S of junction with 167/56,6 May 
1979, Castaner 5667 (MO*), Stone Co.: Ozark National Forest, =) aici DISHES ETON, R11W,S15,NW4 of NW4, 27 
Apr 1993, Hyatt 5251 (MO*).Union Co.: S of El Dorado, at a roadside lcome center 5.7 mi N of Louisiana 
on US Hwy. 167,18 Mar 1995, Hyatt 6159 (MICH**). DELAWARE. INewCastle Co.]: Newcastle, [no date], Canby 
n. (BH*). DISTRICT OF COLUMBIA. [Washington Co.]: Brookland, 14 May 1899, Holm s.n.(MO*);Washington, 
ne date], Vasey 187 (BH**). FLORIDA. Gadsden Co.: near Flat Creek, general vicinity of Sycamore, 6 Mar 1977 
Godfrey 75723 (BRIT**); N of FL Route 20 between Ochlockonee River and Hosford, 15 Mar 1975, Godfrey 74186 
(BRIT**). Liberty Co.: ca.6 mi NE of Bristol, N of Route 12 just W of its junction with Route 271, 14 Mar 1994, Naczi 
3620 (BRIT**). GEORGIA. Clarke Co.: near Winterville, 2 Mar 1929, Miller & Maguire 297 (BH**); GA State College 
Agricultural Farm, 9 Mar 1929, Miller & Maguire 298 (BH**); Princeton Woods, § side of Oconee River, 6 Apr 1929, 
Miller & Maguire 300 (BH). Dade Co.: E side of Route 136, 3.55 mi by road NW of junction with Route 189, 20 Apr 
2004, Werier 2067 (BH**). Lincoln Co.: ner sum Gio Mountain, 5 Apr 1939, Clausen & Trapido 3794 (BH*). 
Rabun Co.: SE of Glade Mountain, 17 Apr | 33 (BH**). Upson Co.: Dripping Rocks, 30 Mar 
1948, Smith et al. 3667 (BH**). ILLINOIS. Al ee Shawnee National Forest, Jonesboro Ranger District, 
Compartment J-104, 5 May 1994, Phillippe 23977 ee Union Co.: Ozark Hill Prairie cane Natural Area, 
Shawnee National Forest, 8 Apr 1992, Phillippe 19778 (MICH**). KENTUCKY. Reale Co.: along Hwy.411,0.4 mi 
E from Hwy. 85, 27 Apr 1987, McKinney 2473 (BRIT*). Lawrence Co.: above state Route | ae ae 0.8 mi NE of 
eek of Benbow Road, 12 Apr 1990, Cusick 28744 (BRIT*). LOUISIANA. te Parish: State Line Road 14.8 mi 
of junction with Hwy. 171,13 Apr 1986, Rettig 1457 (MO**). Claiborne Parish: Kisatchie National Forest Caney 
ee district, 10 Apr 1996, Hyatt 6876 (MICH**). Grant Parish: Kisatchie National Forest Catahoula Ranger 
district, 6 Mar 1995, Hyatt 6157 (MICH**): Kisatchie National Forest, about 3 mi NE of Williana, 3 Mar 1997, Hyatt 
7865 (MO**); Dry Prong community vicinity, 22 Apr 1996, Hyatt 6857 (MICH**). Morehouse Parish: S of LA 835 
by Bryant Main Line Road then E to end of Kitpatrick Road, 30 Apr 1992, Bryson 11441 (MICH**). Natchitoches 
Parish: Kisachie National Forest,ca. 1 mi SE of Goldonna at Saline Bayou on LA Hwy. 156,24 Feb 1998, Hyatt 7860 
(MO**); Kisatchie National Forest, Kisatchie Ranger district, compartment 4, 10 Mar 1998, Hyatt 7872 (MICH**). 


Ouachita Parish: 3 mi SW Cheniere, 11 Apr 1959, Kral 8510 (BRIT**). Rapides Parish: Brushy Creek Riparian Site, 
23 Mar 1998, a ee eee Sabine Parish: LA 175 at Pleasant Hill, 3 Oct 1967, Thomas 5635 (BRIT**). 
Union Parish: y. 2, mi E of H 33/15 in Farmerville, 11 Apr 1986, Rettig 1456 (BRIT**). MARYLAND. 


Baltimore Co.: ieee 7 mi NE ie Baltimore, 5 May 1910, Churchill s.n. (BH**), Caroline Co.: Tuckahoe 
State Park, W of Cherry Lane, 1.2 mi S from junction with Mason Branch Road, 2 May 2004, Werier 2177 (BH*), 
Werier 2179 (BH**).Montgomery Co.: Great Falls, 30 May 1917, ee 1302 (BH**). Prince George Co.: near 
Congress Heights, 25 May 1915, VanEseltine 385 (BH**). MISSISSIP tala Co.: 5.5 mi SSE of McCool, 12 Apr 

1986, Bryson 4152 (MO**); 2.6 mi SSE of Attala-Carroll County line fa MS Hwy. 35,12 Apr 1986, Bryson 4138 
(MO**).Holmes Co.: Holmes County State Park, 16 Mar 1968, McDaniel 10376 (BRIT**). Kemper Co.: White Horse 
Mountain, ca. 10.5 air mi N of Dekalb, 6 Apr 1972,McDaniel 15865 (BRIT**). Lafayette Co.: 4.2 mi S of intersection 
of MS Hwys. 7 and 6 bypass,s of Oxford,9 Apr 1987, Bryson 5327 (MO*). Leake Co.: near Yockanooka ny Creek, ca. 


WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1069 


8 mi WSW Carthage, 2 Apr 1972, McDaniel 15819 (BRIT**). Oktibbeha Co.: 0.5 mi NW of Sturgis, Hannah Hill, 20 

r 1989, Bryson 8463 (MO*).Winston Co.: W Betheden Lutheran Church, 6 Mar 1974, Bryson 318 (BRIT**); 2.5 mi 
NE Louisville,6 Mar 1974, Bryson 317 (BRIT**). MISSOURI. Butler Co.: Poplar Bluff Bottomland Hardwood Natural 
Area, 24 Apr 1982, Castaner 6749 (MO*). Callaway Co.: 0.25 mi E of Williamsville, 25 Apr 1997, Summers & Vander- 
bilt 8062 (MO**). Carter Co.: Mark Twain National Forest, 3 mi S of Van Buren off Hwy. 103 in Spring Valley, 6 May 
1995, Summers & Wallace 7334 (MO*). Laclede Co.: Mark Twain National Forest, Gasconade River, 2 mi W of 
Adam Ford, 24 Jun 1995, Summers 7460 (MO**).Madison Co.: 8 mi S of Twelve Mile Creek, along Hwy.67, 26 Apr 
1987, Castaner 9602 (MO*). Shannon Co.: Middle Hollow, 7 May 1997, Brant 3687 (MO*). Ste. Genevieve Co.: 
Hickory Canyon Natural Area, ca. 2 mi N of Sprott on Hwy. EE, 23 May 1996, Summers et al. 7800 (MO**); Pickle 
Springs Natural Area, ca.0.25 mi E of Hwy. AA, ca. 1.5 mi SE of Hwy. 32, 20 Apr 1996, Summers et al. 7695 (MO**). 
Wayne Co.: Mark Twain National Forest, 2 Apr 1998, Brant 3882 (MO*). NEW JERSEY. Somerset Co.: Chimney 
Rock, Bound Brook, May 1935,Edwards s.n. (BH**). [Gloucester Co.]: Newfield, 20 May 1873, Ellis s.n. (BH**). NEW 
YORK. Orange Co.: Town of Warwick, SE end of Bellvale Mountain, N of village of Greenwood Lake, SW of Route 
17A, 7 May 2003, Werier 1506 and Barbour (BH*). NORTH CAROLINA. Davidson Co.: a little E of Tuckertown 
Road, 1.5 mi by road S of Route 49, 14 Apr 2004, Werier 1992 (BH**), Werier 1993 (BH*), Werier 1994 (BH**); S of 
Route 49,0.6 mi E of bridge over Yadkin River, 14 Apr 2004, Werier 1996 (BH**). Montgomery Co.: Flint Hill Road, 
ca. 2.5 mi by road W of Lovejoy Road, 13 Apr 2004, Werier 1986 (BH**). Swain Co.: 100 yards inside Graham-Swain 
County line on US 129, 11 Apr 1983, Reznicek 7132 (MO*); SE of Route 19 at junction of Route 28, 26 Apr 2004, 
Werier 2128 (BH**). Transylvania Co.: N side of Route 64 just E of Lake Toxaway, 18 Apr 2004, Werier 2044 (BH™*). 
OHIO. Gallia Co.: Symmes Creek Road, 1.25 mi S of Gallia-Centerpoint Road, 9 Apr 1986, Cusick 25054 (MICH*™). 
Jackson Co.: Camp Canter's Cave, N of Caves Road, 16 Apr 1990, Cusick 28752 (MICH**). Scioto Co.: N side of 
Dark Lick Road ce mi Ne oi Perici Road, 19 Apr 1990, McCormack 2094 (MICH**). OKLAHOMA, nate Co.: 


3miWonSH63 klal li Apr 1992 Magrath 1 18592 ee “ side OK Sees Imi 
SE of Little ie eee at Honobia, 19 py 1993, Rezn oe 0353 mer oe ‘ord Co.: 0.75 mi WSW of 
Breezewood, 16 May 1946,Berkheimer7110(BRIT*™).B eee ne = 


3 May 2004, ney 185 (BH**). [Carbon Co.]: ca. 1/8 mi W of Lehigh Gap LV.R R station, 22 May 1927, Pretz 12925 (BH™), 
Centre Co.: Ingleby, 2 mi E of Coburn, 5 May 1936, [Wahl] 2486 (BH*™*); ee 6 a) ae eaten & Wale 2528 (BH**); 
abandoned railroad at Ingleby, 7 May 1977, Keener 3401 (MO**). Huntingdo 29 May 1979, 
Lahham & Keener 15 (BRIT*). Potter Co.: Town of Os N of US Route 6,ca.0.5 mi NW of Galeton, May 2005, Werier 2546 
(BH*). SOUTH CAROLINA. Anderson Co.: 2 mi SE of S Springs, 6 Apr 1939 ee - (BH**). Chesterfield 
Co.: N side of Route 9,W of Thompson Creek,W of Zoar Road, 15 Apr 2004, Werier 2003 (BH**). Darlington Co.: Society Hill, 
[no date], Curtis 106(BH**).[Darlington Co.]: W of Hartsville,9 Apr 1921,Norton C21 (BH), Norton s.n.(BH**).Greenville Co.: 20 
meters SE from Route 414, 2.1 mi by road NE from Route 276, 18 Apr 2004, Werier 2028 (BH*). Lancaster Co.: E of Route 601, 
S of Flat Creek, 16 Apr 2004, Werier 2011 (BH**);E of Great Falls and a little bit E of the Catawba River, 17 Apr 2004, Werier 2024 
(BH**). Oconee Co.: SE side of Stump House Mountain, 7 Apr 1939, Clausen & Trapido 3811 (BH**). Pickens Co.: just N 
Route 288, 1.2 mi by road E of Pumpkintown and Route 8, 18 Apr 2004, Werier 2031 (BH**), Werier 2034 (BH**), Werier fe 
(BH**); just E of SSR 69,0.1 mi by fea N from Route 288, 2.5 mi by road E from Route 178, 18 Apr 2004, Werier 2039 (BH*). 
TENNESSEE. Davidson Co.: Dyer Observatory, 30 Mar 1976, Kral 57466 (MO*).Grundy Co.: NW of state Route 50, 5.7 mi by 
road SW from junction with Route 108,22 eee Werier 2093 (BH**).Polk Co.:N of Route 64,11.1 mi by road E of junction 
a Route 314,20 Apr 2004, Werier 2059 (BH**). Sevier Co.: N of Little Cove Creek Road,4.8 mi by road E of Wear Valley Road, 
Apr 2004, Werier 2134 (BH**). TEXAS. Cass Co.: Hwy. 155, 1.5 mi SW of its junction with Hwy. 59 in Linden, 14 Apr 1 
At 1469 (MO**);3 mi SE Hughes Springs, 2 Apr 1979,Kral 63243 and Carter (BRIT**). Houston Co.: FR 227,0.1 mi NW on 
junction with Davy Crockett Road (NF 526), 10 Apr 1990, 1309 and Wipff (MO**).Smith Co.: Tyler State Park N of Tyler, 
13 Apr 1986, Rettig 1465 (BRIT**). Upshur Co.: Hwy.155,9.5 mi “SW of its junction with Hwy.259 in Cedar Springs, 14 Apr 1986, 
Rettig 1467 (MO**). VIRGINIA. Botetourt Co.: NW of Route 615,0.7 mi by road N of junction with Craig Co.line, 29 Apr 2004 
Werier 2148 (BH**). Brunswick Co.: 7.7 to 7.8 mi by road N of Route 712 from Route 58 in Edgerton, 11 Apr 2004, Werier 1969 
BH**), Werier 1971 (BH**), Werier 1975 (BH*); Route 670, 2.2 to 2.3 mi by road N of Route 46, 11 Apr 2004, Werier 1981 (BH*™*). 
Caroline Co.: near North Anna River, 10 Apr 2004, Werier 1952 (BH), Werier 1957 (BH**), Werier 1959 (BH**).Craig Co.: Potts 
Mountain ca.6 mi NW Castle, 18 May 1983, Wieboldt 4657 (MO*). Powhatan Co.: Powhatan State Wildlife Manage- 
t Area, NW of M 11 Apr 2004, Werier 1963 (BH**). WEST VIRGINIA. Binge Go: county Bette 2, ] TSS — 
Route 2 at Sevan 17 Apr] 991, Cusick 29399 (MICH**). Pleasants Co.: a! f Micki land Creeks, 10 
1984, Cusick 23333 and Ortt Ce aie oo county Route 8, 1 1/2 mi S of US Route 50,8 May 1985, chee 
aes (BRIT*). Tyler Co.: j y Fairview Church, 3 May 1989,Cusick 28021 and Ortt (BRIT**). 


= 
= 


ACKNOWLEDGMENTS 


I thank Anton Reznicek for advice, thoughts, comments, and help on various aspects of 


1070 BRIT.ORG/SIDA 22(2) 


my research project; Susan Reznicek for drawing the illustration; Anita Barbour, James 
(Spider) Barbour, Arthur Haines, and Thomas Wieboldt for assistance with field work: 
Nat Cleavitt for help with analysis of data; Neil Golder for help translating my abstract 
into Spanish; Guy Nesom and Thomas Vining for assistance with the Latin diagnosis; 
Peter Ball, Nat Cleavitt, Bob Dirig, Philip E. Hyatt, Richard LeBlond, Robert Naczi, Anton 
Reznicek, and Troy Weldy for reviewing my manuscript and providing valuable sugges- 
tions; the Bailey Hortorium and its staff for providing space to do my research; and Bob 
Dirig of BH for processing all of my loan requests and helping with other details related 
to my work at BH. Lalso thank the curators from B, BRIT, DOV, GH, MICH, MO, NCU, NY, 
NYS, PH, US, and VPI for loans of specimens and/or assistance during my visits and with 
my inquiries; and Philip E. Hyatt and Robert Naczi for sending recently collected speci- 
mens. The New York State Museum Biological Survey provided funding for survey work 
in the Hudson Highlands of New York which led to the initial discovery of C. reznicekii. 


REFERENCES 


Crins, W.J.and J.H. Rettic. 2002. Carex Linnaeus sect. Acrocystis Dumortier. In: Flora of North America 
Editorial Committee, eds., Flora of North America North of Mexico, Vol. 23. Magnoliophyta: 
Commelinidae (in part): Cyperaceae. Oxford Univ. Press, New York. Pp. 532-545, 

Cusick, A.W. 1992. Carex section Acrocystis (Cyperaceae) in Ohio. Michigan Bot. 31:99-108. 

Fernaco, M.L. 1902. Contributions from the Gray herbarium of Harvard University. The variation of 
some boreal carices. Proc. Amer. Acad. Arts 37:495-513. 

Fernaco, M.L. 1950. Gray's manual of botany. 8th ed. American Book Co., New York. 

KUKENTHAL, G. 1909. Cyperaceae-Caricoideae. In: A. Engler, ed., Das Pflanzenreich. IV. Vol. 20, Heft 38: 
Leipzig: Wilhelm Engelmann. Pp. 1-824. 

LEBLOND, R.J.,A.S. Weakey, A.A. Reznices, and W.J.Crins. 1994. Carex lutea (Cyperaceae), a rare new coastal 
plain endemic from North Carolina. Sida 16:153-161. 

Mackenzie, K.K. 1913. Notes on Carex VII. Carex umbellata and its allies. Bull. Torrey Bot. Club 40:529- 
554. 

Mackenzie, K.K. 1935. Montanae. N.Amer.Fl. 18:185—206. 

McCune, B.and J.B. Gract. 2002. Analysis of ecological communities. MJM Software Design, Gleneden 
Beach, OR. 

Naczi,R.F.C.,R. Krat, and C.T. Bryson. 2001. Carex cumberlandensis,a new species of section Careyanae 
(Cyperaceae) from the eastern United States of America. Sida 19:993-1014. 

Reznicek, A.A. and K. Cameteeke. 1996. Carex porrecta (Cyperaceae), a distinctive new species for north- 
ern South America and Costa Rica. Novon 6:423-425, 

Reznicek, A.A. and P.M. Catuna. 1982. Cyperaceae new to Canada from Long Point, Norfolk County, 
Ontario. Canad. Field-Naturalist 96:184-188. 

Roatson, E.H., J.T. CoLumpus, and E.A. Friar. 2001. Phylogenetic relationships in Cariceae (Cyperaceae) 
based on ITS (nrDNA) and trnT-L-F (cpDNA) region sequences: Assessment of subgeneric and 
sectional relationships in Carex with emphasis on section Acrocystis. Syst. Bot. 26:318-341. 

Roatson, E.H. and E.A. Friar. 2004. Phylogenetic relationships and biogeographic patterns in North 
American members of Carex section Acrocystis (Cyperaceae) using nrDNA ITS and ETS sequence 
data. Pl. Syst. Evol. 243:175-187. 

Tucker, G.C. 1995. Preliminary keys to Carex (Cyperaceae) in New York State. New York Fl. Assoc. Newsl. 
6(2):3-18. 


A NEW SPECIES OF EUGENIA (MYRTACEAE) FROM THE 
MONTEVERDE REGION, COSTA RICA 


Fred R. Barrie 
Missourl Botanical Garden 
PO. Box 299, St. Louis, Missouri 63166, U.S.A. 
ang 


fe; IdM 
artment rela 


1400 5, lake Shore Drive, cee lin inois 60605 
fbarrie@fieldmuseum.org 


ABSTRACT 


on oe | -R eT | \ 1c gee VSR co eer | ] Puntarenas 


Anew species, E 


provinces of Costa Rica. It may be distinguished from E. rhombea Dy chatacters of the leaves and flowers, espe- 
cially the calyx. The species is apparently restricted to cliff edg 1100 to 1350 meters. 


ABSTRACT 


to (AA y 


Se describe una nueva especie, Eugenia haberi B de las provincias de Guanacaste y Puntarenas 
en el lado pacifico de Costa Rica. Puede diferenciarse de E. rhombea por caracteres de las hojas y flores, 
especialmente el caliz. La especie esta restringida aparentemente a bordes de acantilados a elevaciones de 1100 a 


1350 metros. 


INTRODUCTION 


With about 700 species, Eugenia is the largest of the New World genera of Myrtaceae. 
Forty-nine Eugenia are currently known to occur in Costa Rica, including 21 recently 
described (Barrie 2005) and the species described here. Eugenia is characterized by a race- 
mose or fasciculate inflorescence, flowers with an open, four-lobed calyx and an embryo 
that is an undifferentiated mass of tissue. 


reco haberi Barrie, sp. nov. (Fig. 1). Type: COSTA RICA: PUNTARENAS. Monteverde, Eston’s Cliff Edge, 
350 m, | Jan 1979 (£1, fr), W. Haber 263 (HOLOTYPE: MO}; ISOTYPES: MEXU!, MO!, N 


lint? 
oO r 


, pedicellis brevioribus et lobis 


Eugenia rhombea (O. Berg) Krug et Urb. aemulans, differt foli 
calyce minoribus. 

Small tree; young growth glabrous or with a few hairs on the margins of the bracts, 
bracteoles and calyx. Branchlets compressed at the outer nodes; bark reddish brown. 
Leaves narrowly elliptic, the blades 3-8 x 1-2.5 cm, 2-5 times as long as wide, coriaceous, 
drying a concolorous green or somewhat darker above, the midvein and margins tan; 
midvein convex on both surfaces, often with a narrow ridge above: lateral veins 15-20 
per side, straight, often obscure above; marginal veins similar to the laterals, straight, ca. 
1mm from the margins; glands numerous on both surfaces, sometimes punctuate above: 
base narrowly cuneate to weakly concave; margins cartilaginous, decurrent along the 
petiole; apex acuminate to caudate-acuminate, the tip ultimately rounded; petioles 2-5 
mm long, flattened or weakly concave dorsally. Inflorescences axillary, racemose, soli- 
tary; axis 1-5 x 1 mm long; flowers 4-10; bracts 0.5 mm in diameter, persistent, circular, 
the margins scarious, with or without a few hairs; buds 2-3 mm long, ovoid. Flowers pedi- 
cellate, the pedicels 4-12 mm long, with prominent pellucid glands; bracteoles 0.7-1 
0.5 mm, persistent at least through anthesis, ovate, prominently glandular, the base free 


SIDA 22(2): 1071- 1073. 2006 


1072 BRIT.ORG/SIDA 22(2) 


MISSOURI 
BOTANICAL GARDEN 
HERBARIUM 


N° 2740686 


HOLOTYPE of Eugenia haberi Barrie 


F.R, Barrie 2006 
Mi 


COSTA RICA 
Puntarenas 


Family: MYRTACEAE 
Eugena ct, COStariCensit kery 
Monteverde, Eston's cliffedge, 1350 m. 
Small tree. 
W. Haber 263 2 January 1979. 
MISCANID! BATAMIFAL ‘RA OMeM UeRR anne cans 


Fic. 1. Holotype of Eugeni ( , MO) 


BARRIE, A NEW SPECIES OF EUGENIA FROM COSTA RICA 1073 


or slightly connate, the margins entire or scarious, with or without a few hairs; hy- 
panthium 1.5-2 mm long, campanulate; calyx lobes in subequal pairs, 0.7-1 x 0.5 mm, 
elliptic, prominently glandular, the margins entire or witha few hairs, the apex rounded: 
petals ca. + x 3.5 mm, ovate, the margins with or without a few hairs, the apex rounded; 
disk 1.2-2 mm in diameter, glabrous; stamens ca. 50, 3-5 mm long; style 5-6 mm long, 
glabrous. Fruits 6-11 x 6-10 mm, globose or obovoid, mature color black; pericarp thin- 
walled, glabrous; calyx persistent but reduced. 

Eugenia haberi is a small tree found along cliff edges on the Pacific slope in 
Guanacaste and Puntarenas provinces at elevations of 1100-1350 meters. In many respect 
it resembles E. rhombea (O. Berg) Krug & Urb., a species of coastal areas with a circum- 
Caribbean distribution. Although E. rhombea occurs in both Panama and Nicaragua, it 
has yet to be found in Costa Rica. The similarities between the two species include gen- 
eral habit, the overall absence of vestiture, with hairs restricted to the margins of the bracts, 
bracteoles and perianth, and the coriaceous leaves with cartilaginous margins. The two 
may be differentiated by the shape of the leaves, which in E. haberi is narrowly elliptic 
with a tapering, narrowly cuneate base and an acuminate to caudate-acuminate apex. In 
E. rhombea, the leaves are ovate to elliptic with a rounded or broadly cuneate base and an 
acute apex. The inflorescences in E. rhombed are often superficially fasciculate, and the 
pedicels of the flowers and fruits are up to 30 mm long. In E. haberi the inflorescence 
rachis may be short but it is apparent and the pedicels do not exceed 12 mm. The calyx 
lobes of E. rhombea are much larger and more prominent, up to 4 mm long and 2 mm 
wide, and are prominent in flower and fruit, retaining a pale color that contrasts with the 
dark body of the mature fruit. The calyx persists but is not conspicuous on the fruits of E. 
haberi. 

Etymology.—The species is na med for Bill Haber, the collector of the type, whose deep 
and extensive knowledge of the flora and fauna of the Monteverde region is exceeded 
ae by his Best in sharing the information with others. 


Addi Ico] |. COSTA RICA. Guanacaste: Canton Liberia, Parque Nacional Guanacaste, ee 

Cacao, 1100 m, 10°55'45'N, 85°28'15"W, 22 Nov 1990 (v?) C. Chavez 391 (NB); Estacion Cacao, 7 May 1 
Chavez 553 (CR, E INB, MO); Estacion Cacao, 11 Apr 1991 (fr), C. Chavez 541(CR, INB, MO). Sania ner 
Hoge Uppan, 10 Feb 1979 (fl) W.A. Haber 263 (CR); Monteverde, lower community on Pacific slope, 1350 m, 10°18'N, 
84°48'W, 5 Jul 1991 (fr) W. Haber & C. Ivey 10720 (CR, F, INB). 


ACKNOWLEDGMENTS 
This research was conducted while in residence in the Botany Department of the Field 
Museum. I thank the department and the chair, Michael O. Dillon, for their generous, con- 
tinuing support. Thanks to Bill Haber for sharing his knowledge of the field characteris- 
tics of this species, and to Bruce Holst, Lucia Kawasaki, and Les Landrum for reviewing 
this manuscript. 


REFERENCES 


Barrie, FR. 2005. Thirty-five new species of Eugenia (Myrtaceae) from Mesoamerica. Novon 15:4-49. 


1074 BRIT.ORG/SIDA 22(2) 


Book NOTICE 


Neat K. VAN ALFREN, GEORGE BRUENING, and WILLIAM O. Dawson (eds). 2006. Annual Review 
of Phytopathology: Volume 44, 2006. (ISBN 0-8243-1344-5, hbk; ISSN 0066-4286). 
Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, 
US.A. (Orders: ane awe org, 800-523-8635, 650-493-4400, 650-424-0910 
fax: onlineactivati rg). $188.00 (USA), 538 pp., 7 3/8" x 91/4" 


Annual Review of Phytopathology: 


mH 


Contents of Volume 44 of 
A Retrospective of an Unconventionally Trained Plant Pathologist: Plant Diseases to Molecular Plant Pathology 
The Current and Future Dynamics of Disease in Plant Communities 

A Catalogue of rhe Effector Secretome of Plant Pathogenic Oomycetes 
Genome Packaging by Spherical Plant RNA Viruses 

Quantification ie Modeling of Crop Losses: A Review of Purposes 
Nonsystemic Bunt Fungi—Tilletia indica and T. horrida: A Review of H 
Significance of Inducible Defense-related Proteins in Infected Plants 
Coexistence of Related Pathogen Species on Arable Crops in Space and Time 

ng Nonpersistent and Semipersistent Transmission of Plant Viruses 


istory, Systematics, and Biology 


Virus-Vector Interactions Mediatir 
Breeding for Disease Resistance in the Major Cool-Season Turf grasses 

Molecular Ecology and Emergence of Tropical Plant Viruses 

Biology of Flower-Infecting Fungi 

A Model Plant Pathogen from the Kingdom Animalia: Heterodera glycines, the Soybean Cyst 
Nematode 

Reveals What a An Enterobacterial Plant Pathogen 


Comparative yeno 

The Dawn of Fungal Pathogen Gen 

Fitness of Human Enteric = ee on nPl ants and Implications for Food Safety 

The Role of Ethylene in Host-Pathogen Interactions 

mae Cc ae ae in Fluorescent Pseudomonas spp. Biosynthesis and Regulation 
Long-Distance RNA-RNA Interactions in Plant Virus Gene Expression and Replication 

Evolution of Plant ca ie in Streptomyces 

Climate Change Effects on Plant Disease: Genomes to Ecosystems 


SIDA 22(2): 1074. 2006 


TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS 
(ASTERACEAE: ASTEREAE) 


Guy L.Nesom 
Botanical Research Institute of Texas 
509 Pecan Street 
Fort Worth, Texas 76102-4060, U.S.A. 


ABSTRACT 


The eastern North American Symp} ichum patens has been treated by Jones (1983, 1992) and other olan 
to include var. patens, var. patentissimum, and var. gracile. These taxa h ily by habit, 
involucral size and vestiture, and phyllary orientation. aa cisHine Hons between v var. patens and var. 
gracile have been sedaiee as or APE ne and nS two ie a have b wide region; 


the present study h them inan Sn or evolutionarily mean- 


ingful way, and var. gracile is not fecconiecd Variety patens sensu lato appr eney Comprises two open on 


three or four tetraploid population systems. Variety } 
tetraploid, and it ene ae ae eae var. paren: ai their oe are mera or peaks A 
county-level map shows th the two infraspecific taxa a S ee as interpreted in the present 
| Loi previous studies. 


Ana) Alt +1 “ cr BE aaron ‘ sak ack 


study. A similar 


RESUMEN 


eal patens del este de Norte América ha sido tratado por Jones (1983, 1992) y otros botanicos para 
incluir las var. patens, var. patentissimum, y var. gracile. Bae taxa han sido esters panels mente por el 


ae tamano involucral y pubescencia, y la orientacion de los filarios. I g var. 
patens y var. gracile ha sido vista como solapada y los dos ta ido i d impa u 
region ae el presente estudio concluye que no es ae distinguirlos ae un moso que no sea Oranere oen 


una linea evolutiva significativa, y la var. gracile no se reconoce. La vaneudG ae sensu lato comprende 
aparentemente dos sistemas de poblaciones diploides y tres 0 cuatro tetraploides. La variedad patentissimum es 
ead lanes diferente ys constantemente tetraploide, y se intergrada con la var. ci es es ToT IOIGS, 


alli do ua 1 Un mapa a nivel de condado muestra la dist de los dos 


taxa eee de S. ee tal como se secre en a Diesenis estudio. un mapa similar muestra la 


distribucion ploidal de la especie, 


The eastern North American Symphyotrichum patens (Aiton) G.L.Nesom has been treated 
(especially in detail by R.L. Jones 1983, 1992, as Aster patens) to include var. patens, var. 
gracile (Hook.) G.L.Nesom, and var. patentissimum (Lindl.) G.L.Nesom. Infraspecific taxa 
have been distinguished primarily by habit, involucral size and vestiture, and phyllary 
orientation. Morphological differences, however, especially between var. patens and var. 
gracile, have been construed as overlapping and those taxa have been viewed as broadly 
sympatric. 

Differences among the varieties have been characterized as in the key below, which 
is amalgamated mostly from contrasts and comments by Cronquist (1980), A.G. Jones 
(1992), Jones (1983), and Brouillet et al. (2006). 

1. Involucres broadly turbinate, 8-12 mm oa phyllaries often in 5-7 series, appressed, obtu 

mid phyllaries ovate-lanceolate, 1.2-1.7 mm wide, densely strigillose to sericeous- pene 

eglandular to sparsely and minutely a aeanae var. patentissimum 
Involucres campanulate or slenderly turbinate, 5.0-7.5 mm long; phyllaries in 4-5(-6 ee 

often at least somewhat squarrose, acute to acuminate, mid phyllaries linear-lanceolate, 0.7—1 

mm wide, sparsely to densely strigillose, sparsely 2 oo sessile- or stipitate- glandular. 


2. Plants slender branches long conspicuous! 1d minutely bracteate; princ | cauline leaves 


—_— 
aii 


SIDA 22(2): 1075- 1080. 2006 


1076 BRIT.ORG/SIDA 22(2) 


1-3(-4) cm long, 0.5-1(-1.5) cm wide; involucres 5.5-6.5(-7.5) mm long; phyllaries sparsely 

glandular and densely strigillose, mid 0.7—1 mm wide var. gracile 
2. Plants more robust, branches shorter, with fewer and larger bracts; principal cauline leaves 

3-6(-8) cm long, 1-2 cm wide; involucres 6-7.5 mm long; phyllaries densely glandular and 

sparsely strigillose, mid 1—1.2 mm wide var. patens 


Variety patentissimum occupies a geographic range in the Interior Uplands west of the 
Mississippi River, except for populations in southern Illinois. Jones (1983) mapped scat- 
tered collections of var. patentissimum from Texas and Louisiana; Cronquist (1980) included 
Mississippi in the range. Variety patens has been recorded from Maine (Magee & Ahles 
1999), New Hampshire, and Masschusetts south to Florida, thence westward to Texas, Okla- 
homa, and Kansas (Jones 1983). Variety gracile was recorded by Jones (1983) in Texas, Okla- 
homa, Kansas, Missouri, Arkansas, Louisiana, Mississippi, and Alabama — sympatric in 
each state with var. patens and sympatric in Missouri, Arkansas, and Oklahoma with var. 
patentissimus. A.G. Jones (1992) viewed all three taxa as present in Oklahoma. Brouillet et 
al. (2006, Flora of North America treatment) presented an expanded geographic concept 
of var. gracile, following Cronquist (1980), both of whom recognized its occurrence in 
Florida, Georgia, South Carolina, North Carolina, Virginia, Tennessee, and Maryland, far 
into the range of var. patens. In the view of both R.L. Jones and of Brouillet et al, the range of 
var. gracile is completely sympatric with that of var. patens. Variety patens in the concept 
of Brouillet et al. occurs alone from New Jersey and Delaware northward to Maine and west- 
ward from Pennsylvania and New York to Indiana and Illinois. 

All chromosome number reports of var. patentissimum have been tetraploid (2n = 
20). Many populations of var. patens (sensu Jones 1983) have been reported as tetraploid 
(counts from Massachusetts, Illinois, Indiana, New Jersey, Kentucky, Tennessee, Virginia, 
North Carolina, South Carolina, Georgia, Alabama, and Mississippi); diploids have been 
reported from western Alabama (Jones 1983; Semple 1984). Variety gracile has been re- 
corded at diploid level (2n = 10) in Alabama, Mississippi, Louisiana, Texas,and Oklahoma. 
Tetraploid counts of var. gracile have been reported from Louisiana and Texas Jones 1983; 
Semple 1984). 

Artificial intervarietal crosses between tetraploids (var. patens x var. patentissimum) 
“produced good seed sets, and the hybrid offspring were intermediate in morphology” 
(Jones 1983, p. 60). Crosses between var. patens and Symphyotrichum phlogifolium (both 
tetraploid) were as successful as intervarietal crosses. In contrast, “only a few crosses were 
attempted between diploid and tetraploid S. patens, and none of these were successful” 

OO); 


— 


The concept of Symphyotrichum patens in earlier literature has included S. 
georgianum (Alexander) Nesom and S. phlogifolium (Muhl. ex Willd.) Nesom, but Jones 
(1983) elevated these taxa to specific rank, and most later authors, including Brouillet et 
al. (2006), have followed these concepts, with Cronquist (1980) and Gleason and Cronquist 
(1991) being notable exceptions. Neither S. georgianum nor S. phlogifolium intergrades 
morphologically with other taxa; S. georgianum is a decaploid (2n = 50) while S. 
phlogifolium isa tetraploid (2n = 20), and both occur mostly within the geographic range 
of tetraploid S. patens var. patens. Another distinct but closely related species, S. 
grandifolium (L.) Nesom, is known only as a dodecaploid (2n = 60). The chromosome 
counts for these three taxa were first reported by R.L. Jones and A.G. Jones (1979). 

The present study reexamines the taxonomic distinctions and geographic distribu- 
tions of the infraspecific taxa of Symphyotrichum patens. The study is primarily based 


NESOM, TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS 1077 


on specimens examined from BRITSMU, MISS, MO, NCU, TEX-LL, and VDB (including 
many of the collections made by R.L. Jones in his Vanderbilt dissertation study). The col- 
lection localities of these are mapped in Figure 1. 

Semple (1984) mapped chromosome number reports for Symphyotrichum patens, as 
similarly done here, but the present map (Fig. 2) is more precisely represented and the 
modified taxonomy gives it a different perspective. Chromosome counts have been made 
by various botanists but mostly by Semple and Brouillet (1980), Jones (1983), and Semple 
(1984), as summarized by Semple (1984), with an addition by Jones and Smogor (1984). 
Several chromosome counts, vouchered by collections by John Semple in 1991 and depos- 
ited at TEX and MO, apparently have not been reported in literature: Kentucky, Whitley 
Co., 2n = 20 (MO); North Carolina: Wilkes Co., 2n = 20 (MO); Ohio: Scioto Co., 2n = 20 
(MO); Texas: Harrison Co., 2n = 10 (TEX) and Fort Bend Co., 2n = 20 (TEX), West Virginia: 
Pendleton Co., 2n = 20 (MO). 


RESULTS AND DISCUSSION 


Observations of Jones (1983) and of Brouillet et al. (2006) are confirmed here: plants re- 
ferable to var. patens (using criteria as in the key above) occur essentially throughout the 
range of var. gracile, and plants referable to var. gracile occur through part of the range of 
var. patens. Map 14A of Jones (1983) indicates that the eastern extension of var. gracile is 
abruptly demarcated along the Mississippi-Alabama border, but the present study has 
not confirmed this. Typical var. gracile-like plants occur northeastward at least into North 
Carolina. Similarly, collections from Texas and Louisiana resemble plants identified as 
var. patens in Atlantic states. There appears to be a weak east-to-west trend in increase of 
nonglandular involucral vestiture and, beginning in Alabama westward, development 
of longer and more bracteate branches, but the present study has not found any geo- 
graphic zone of morphological discontinuity that would enable or justify the recognition 
of two taxa. Continued formal recognition of var. gracile perpetuates a typological defi- 
nition. In the present view, recognition of broadly sympatric varieties (as var. gracile with 
var. patens) would imply either that the two were reproductively isolated (thus each bet- 
ter treated at specific rank) or else that they represent populational variants (thus not 
justifiably treated at varietal rank). 

Variety patentissimum isa distinctive entity. It overlaps in distribution (Fig. 1) with 
tetraploid var. patens, and putative hybrids and introgressants apparently occur in the 
region of sympatry: Arkansas, Oklahoma, the southwest corner of Kansas, and (east of 
the Mississippi River) southern Illinois and extreme western Kentucky (intermediates 
not represented on map). Some measure of reproductive isolation may be active, as typi- 
cal plants of both taxa have been collected in relatively close proximity in their region of 
sympatry. Typical var. patentissimum occurs as far south as Cleburne and Dallas coun- 
ties in south-central Arkansas and Sevier and Little River counties in the southwest cor- 
ner of the state; var. patent issimum and var. patens and inte Li occur in Craighead 
and Poinsett counties near the northeast corner of the state. 

Plants and populations of var. patens sensu lato of different ploidy level do not ap- 
pear to be intermixed. More counts, of course, might reveal that diploids and tetraploids 
occur in sympatry, but based on current evidence, diploid populations within var. patens 
apparently occur in two geographic groups, as hypothesized in Fig. 2. Tetraploids appar- 
ently occur in four groups (or three groups, if tetraploidy has been derived only once on 
the east and west sides of the Mississippi River) (Fig. 2). More counts might reveal that 


1078 BRIT.ORG/SIDA 22(2 


A var. patentissimum 


@ var. patens 


Be 1. eDistabuten i Symphyoricnum patens var. patens and var. patentissimum. Symbols indicate counties from which specimens 

y. The record for var. patens in tay oe Kansas, is s fide R.L. Jones (1992); that for var. patens in 
Portage Co., Ohio, is fide Fisher (1988) Th g | New York, Rhode Island, Connecticut, New 
Hampshire, Massachusetts, and Maine. 


diploids are continously distributed along the outer coastal plain 

The nomenclatural type (lectotype) of var. gracile was collected in Washington Par- 
ish, Louisiana, a region where tetraploids, but not diploids, have been found. Morphologi- 
cal or ecological features that might be diagnostic of these population systems of differ- 
ent ploidy level are not apparent. Diploid and tetraploid plants of var. patens in Mississippi, 
for example, as well as in Texas, show similar ranges of morphological variability. Even 
so, it may be assumed that the ploidal systems within var. patens are isolated from adja- 
cent ones by production of triploid hybrids or some other incompatibility Jones noted 
that experimental 2x-4x crosses were not “successful”. 


ACKNOWLEDGMENTS 
A loan of specimens from MISS was critical for studying the pattern of variability in the 
Gulf Coast region. | also am grateful for help from staff at MO, NCU, and TEX-LL during 
visits there and for helpful reviews by Ron Jones and Luc Brouillet. 


NESOM, TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS 1079 

ge es eel ge os a ried Decree Tes oe 1 Let ie 4 age: 

He FR EP STN ci re Or eke 
eons MELEE Rameas cammMite MCPUIR EE Eor nes Varelit CC@p7 CN 


& 
Zauec eeeeeeneee 


— 


r 
tt 


Ape 
en 
sepeaeey \ 


© var. patens, tetraploid 


@ var. patens, diploid 


Fic? Nictrihuti linc L * oh nw [n* a 


occur in two groups. Tetraploids of var. p I k 1 population sy 


REFERENCES 


Broulttet, L., J.C. Sempte, G.A. ALLEN, and S.D. Sunpeer. 2006. Symphyotrichum (Astereae).In: Flora of North 
America Editorial Committee, eds. Flora of North America north of Mexico. Volume 20. Oxford 
University Press, New York and Oxford. Pp.465—539, 

Cronauist, A. 1980. Vascular flora of the southeastern United States, Vol.|. Asteraceae. Univ. of North 
Carolina Press, Chapel Hill. 

Fisuer, T.R. 1988. The Dicotyledonae of Ohio. Part 3. Asteraceae. Ohio State Univ. Press, Columbus. 

Gteason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and 
adjacent Canada (ed. 2). The New York Botanical Garden, New York. 

Jones, A.G. 1992. Aster and Brachyactis in Oklahoma. Sida, Bot. Misc. 8:1-46. 

Jones, A.G. and R.A. Smocor. 1984. In:IOPB chromosome number reports LXXXII. Taxon 33:1 26-134. 

Jones, R.L.and A.G. Jones. 1979. In: IOPB chromosome reports LXIV. Taxon 28:391-408. 

Jones, R.L. 1983. A systematic study of Aster section Patentes (Asteraceae). Sida 10:41-81. 

Jones, R.L. 1992. Additional studies of Aster georgianus,A. patens, and A. phlogifolius (Asteraceae). Sida 
55305 5.15. 


1080 BRIT.ORG/SIDA 22(2) 


Macee, D.W. and H.E. Atits. 1999. Flora of the Northeast: A manual of the vascular flora of New En- 
gland and adjacent New York. Univ. of Massachusetts Press, Amherst. 

Sempit, J.C. 1984. Cytogeographic studies on North American asters. |. Range surveys of Virgulus 
adnatus, V. concolor, V. georgianus, V. grandiflorus, V.novae-angliae, V. oblongifolius, V. patens, and V. 
walteri. Amer. J. Bot. 71:522-531. 

Semece, J.C. and L. Brouiter. 1980. Chromosome numbers and satellite chromosome morphology in 
Aster and Lasallea. Amer. J. Bot. 67:1027-1039. 


== 


DREPANOSTACHYUM FALCATUM VAR. SENGTEEANUM: 
IDENTITY AND ORIGINS (POACEAE: BAMBUSOIDEAE) 


C.M.A. Stapleton 


Royal Botanic Gardens, Kew 
Richmond, Surrey 
TW9 3AB, UK 
chris_stapleton@onetel.com 


ABSTRACT 


} } } } Leet 5 mies 


Drepanostachyum falcatum var. sengteeanum Stapleton, a variety of | g g I 

Himalayas and cultivated in the west for over a century is discussed. The application and the typification of the 
_ name under which it was once grown, Arundinaria falcata var. glomerata Gamble, are considered, along with the 
application of the name Arundinaria falcata Nees. Arundinaria falcata var. glomerata is lectotypilied in accor- 


dance with the protologue and current use of the name in India, placing it in synonymy of Drepanostachyum 
falcatum (Nees) Keng f,, for which an epitype is designated to support its own incomplete lectotype. 


RESUMEN 


Se discute Drepanostachyum falcatum var. sengteeanum Stapleton, una variedad de bambu que se piensa que se 


origino en el Himalaya y se ha cultivado en el oeste durante mas de un siglo. Se consideran la aplicacion y la 
ape ficacion ose pole palo el ane He cultivado, Ar undinaria aay var. glomerata Gamble, junto con la 
ifica A | ta var. slomenas de sewerdoc conel 

ae Pn . 


t ej 
pro 4] 1 de] } la India. colocandolo en | 


8 A 


LULU 
Keng 7 ee el que se designa un epitipo para apoyar su lectotipo i plet 


A bamboo with a prominent and distinctive ring of hairs around the culm sheath base 
and on the young culm nodes was in cultivation at Kew around 1900, without any record 
of its provenance. It was misidentified (Bean 1907) as Arundinaria falcata Nees var. 
glomerata Gamble, a little-known variety from Uttar Pradesh in the NW Indian Himalaya, 
described on the basis of a very dubious floral characteristic. Arundinaria falcata is 
curently placed in the genus Drepanostachyum Keng f. A bamboo with very similar char- 
acteristics started to flower in California in 1994, and its seed has been widely distrib- 
uted, under different names. 

Ithough initially identified as Himalayacalamus falconeri, this graceful Califor- 
nian bamboo was later distributed as Drepanostachyum glomeratum Hort. based on D. 
falcatum var. glomeratum, and then as Drepanostachyum sengteeanum Hort It is proving 
somewhat hardier than all other bamboos in the genus Drepanostachyum, and is thus of 
considerable horticultural merit. It has recently been described as D. falcatum var. 
sengteeanum (Stapleton 2006), buta detailed nomenclatural treatment is required for the 
application of names, D.falcatum and D. falcatum var. glomeratum, to both wild and cul- 
tivated plants. 


Drepanostachyum falcatum 

When describing Arundinaria falcata, Nees (1834) cited ‘Nepalia’ collections made by 
Royle. Chao and Renvoize (1989) designated a Royle collection labelled ‘NW India’ from 
Hooker’s Herbarium (K) as lectotype of Arundinaria falcata. Although this appears con 
Becton no Royle collection from Nepal is extant, and it is actually quite unlikely that 
Royle or his collectors ever entered Nepal, as they concentrated their collecting activities 


SIDA 22(2): 1081— 1086. 2006 


1082 BRIT.ORG/SIDA 22(2) 


to the north and west of their base in Saharanpur, Uttar Pradesh (Edmondson, pers. 
comm.). The Himalayan passes further to the east were little-known, except to some 17th 
Century Jesuits, because of the ‘jealous policies’ of Chinese and Nepalese powers (Royle 
1839). 

For identification of species of Drepanostachyum, culm sheath characters are criti- 
cal, but the culm sheaths of D.falcatum are not well represented in the original material 
at K, nor in the Roylean Herbarium (LIV). One small, glabrous culm sheath is present in 
the lectotype. Munro (1868), Gamble (1896), Tewari (1993) and Negi and Naithani (1994) 
all record the culm sheath of D. falcatum as glabrous, reflecting the characteristics of a 
large number of later collections, made from Himachal Pradesh to Nepal, with which 
the lectotype collections agree in other characters. Drepanostachyum falcatum in the wild 
is clearly interpreted asa bamboo with glabrous culm sheaths (which actually have sparse, 
inconspicuous, white hairs when young). 

Plants of Drepanostachyum falcatum introduced into western cultivation on differ- 
ent occasions also have essentially glabrous culm sheaths. Stapf (1904) was the first to 
identify these plants correctly. They were initially cultivated in S Europe as Bambusa 
gracilis A.& C. Riviére, while the name A. falcata was misapplied to the hardier species 
now knownas Himalayacalamus falconeri(Munro) Keng f., known then as Arundinaria 
nobilis Mitford. Stapf (1904) concluded that early introductions probably came from Naini 
Tal or Mussoori in Uttar Pradesh around 1840. 

The consensus is clear that the name A. falcata applies to a species found in the 
Himalayas from Himachal Pradesh to Kathmandu, and also found in cultivation in Eu- 
rope, where it was known as Bambusa gracilis. This species has almost completely gla- 
brous culm sheaths, although the culm nodes may initially have sparse, short hairs be- 
low. A specimen from Uttar Pradesh is selected here as epitype, to support the incomplete 
Royle lectotype by indicating culm sheath characteristics. 


Arundinaria falcata var. glomerata Gamble 

Arundinaria falcata var. glomerata Gamble was initially distinguished solely on the ba- 
sis of agglomeration of 3-4 spikelets in close racemes or panicles. This followed an ear- 
lier, identical distinction (Munro 1868) between two unnamed developmental forms of 
A. falcata, listed as var.a and var. b. Munro clearly explained that these forms merely rep- 
resented different stages of inflorescence development. He observed that as the season 
advances branch proliferation leads to compound rather than paniculate forms, with 
shorter racemes of 3-4 spikelets (var. b), totally unlike t earlier on younger shoots 
(var. a). He cited a December collection, Wallich 5035 (K-W), from Chisapong, near 
Kathmandu, Nepal as representative of the later developmental form, var. b. 

It is not clear whether Gamble (1896) followed Munro’ interpretation of the devel- 
opmental nature of this distinction. Gamble’s brief description repeated that of Munro 
for var. b. He indicated in a plate caption that illustrations of var. glomerata were from 
collections made by Bagshawe in 1879. This action was interpreted by Chao and Renvoize 
(1989) as designation of a type for A. falcata var. glomerata Gamble, and they cited a 
Bagshawe collection at K as holotype, albeit a collection made in 1878 not 1879. They 
listed var. glomerata in synonymy of Sinarundinaria falcata. 

Tewari (1993) subsequently gave the variety full recognition, providing a detailed 
description attributed to Pandey ined. This, however, differed little from that of the type 
variety. Culm sheath hairs were not mentioned at all. The description of the culm sheaths 


STAPLETON CENCTEEANIIN 4083 


as shining below probably meant having a glossy adaxial, internal surface, rather than 
being basally glabrous, but there was certainly no mention of any prominent basal ring 
of hairs. Naithani and Chandra (1998) and Seethalakshmi and Kumar (1998) merely in- 
cluded it in synonymy of Drepanostachyum falcatum. 

The name A. falcata var. glomerata has thus been applied to two taxa, one a bamboo 
of unknown origin cultivated at Kew, the other an Indian bamboo from the borders of 
Himachal and Uttar Pradesh. The principal diagnostic character of the cultivated plants 
is a prominent ring of hairs around the base of the culm sheath and the culm node. This 
is not seen convincingly in any of the collections of wild material. Other characteristics 
of the cultivated plants include long sparse hairs on only one side of the base of the leaf 
midrib (costa) rather than both sides, densely scabrous lemmas and paleas, and pubes- 
cence only at the tips of rhachilla segments. These characteristics are also not present in 
the wild material, which therefore cannot be considered the same taxon as the cultivated 
plants. 


Typification of var. glomerata 

Original material consists of three differently labelled collections at K from Gamble’s 
herbarium, all annotated by him as Arundinaria falcata var. glomerata. To complicate 
matters, as is usually the case with older bamboo collections including both fertile and 
sterile material, these collections represent at least two, and possibly three different spe- 
cies, with mixed collections mounted together on the same sheets. 

Only one collection, from Jaunsar Bawa, Bagshawe s.n. in 1879, was literally cited in 
the protologue. The culm nodes and persistent portions of the culm sheath bases are al- 
most completely glabrous. 

A second, different collection includes the specimen actually illustrated by Gamble, 
and this, therefore, can be construed as part of the protologue. This sheet, annotated as 
type by Chao, is simply labelled ‘Comm. Brandis, Jaunsar’. As ‘specimens’ collected by 
Bagshawe’ was the citation this could be a further Bagshawe 1879 collection, sent by 
Brandis. It is clearly mixed, with old, unidentifiable flowers of a Drepanostachyum spe- 
cies as well as leaves of a different species, probably Himalayacalamus falconeri, with 
smooth culms, entire leaf sheath ligules and distinguishable tessellation of veins on some 
of the leaves, which are broader, glabrous, and less cuneate than those of D. falcatum. 

Gamble dissected spikelets from these two collections. His drawings remain attached 
to the Bagshawe 1879 collection, with the recognisable spikelets he illustrated remaining 
in capsules on the two sheets. These drawings were reproduced when describing his var. 
glomerata, but interestingly were used for the plate of the type variety, not that of var. 
glomerata, suggesting that Gamble himself actually had scant regard for any difference 
between the two varieties. 

The third collection from Gamble’s herbarium annotated as var. glomerata, also from 
Jaunsar Bawa, but in 1878, is Bagshawe 6608. This was cited as holotype of A. falcata var. 
glomerata by Chao and Renvoize (1989), and is annotated as type by Renvoize. It again 
has no culm sheaths, but like the second collection, has some short hairs below the culm 
node. 

To summarise ‘specimens collected by C. Bagshawe.,.... in 1879’ was the citation in the 
protologue. A collection labelled thus is at K. A second, mixed collection, sent by Brandis, 
was actually illustrated in the protologue, and annotated as type by Chao, while a third 
collection, Bagshawe 6608 in 1878, a date that conflicts with the protologue, was cited as 


1084 BRIT.ORG/SIDA 22(2) 


‘holotype’ by Chao and Renvoize (1989). The collection cited in the protologue has com- 
pletely glabrous culm nodes, while the other two collections have distinct, though very 
short, hairs below the nodes. 

It seems more appropriate for the collection bearing the details cited in the protologue 
to be taken as holotype, rather than the collection stated as ‘holotype’ by Chao and 
Renvoize (1989). Although that collection was part of the original material, as it was nei- 
ther cited nor illustrated in the protologue, their citation of it as holotype cannot be con- 
sidered an acceptable lectotypification. 

The designation here as lectotype of the only sheet actually labelled Bagshawe, 
Jaunsar Bawa, 1879, is in accordance with both the protologue, and with current applica- 
tion of the name in India (Tewari 1993). As the collection actually illustrated in the 
protologue is recognisable, and citation of Bagshawe, 1879 was merely indicating (possi- 
bly incorrectly) which collection was illustrated, there would appear to be two elements 
to consider as syntypes. 

The name D. falcatum var. glomeratum Gamble is misleading, and seems merely to 
representa later stage of inflorescence development, as Munro originally intended (1868). 
There is indeed no separate variety of D.falcatum with agglomerated spikelets. It might be 
considered more appropriate for Gamble to have followed the typification of Munro’s ear- 
lier unnamed variety. However, Wallich 5035 is one of the worst bamboo specimens ever 
preserved in a herbarium, being so decrepit that it has practically no spikelets left at al 

Drepanostachyum falcatum var. glomeratum, as interpreted and described in Tewari 
(1993) and lectotypified here according to the protologue, differs little if at all from the 
type variety. The precise origin of this type is not known, but it is probably from the same 
district as the epitype of D. falcatum designated here. Culm sheaths are not known, but 
basal portions remaining attached to the culm nodes of the lectotype are completely gla- 
brous. 


— 


ee falcatum (Nees) Keng f., J. Bamboo Res. 2:16. 1983. Types: INDIA:‘N.W India, 
Royle (LECTOTYPE: K, anne in Chao and Renvoize 1989); INDIA. UTTAR PRADESH: Chakrata, Sep 1898, 
Gamble 27256 (epityy gnated here: kK) 


Sides Nees, Linnaea 9:478. 1834. 

Chimonobam busa falcata (Nees) Nakai in, J. ce Arbor. 6:151. 1925. 

Fargesia falcata (Nees) TP. Yi, Fl. Xizangica 5:33. 19% 

Sinarundinaria falcata (Nees) CS. Chao & ent €, ieee Bull. 44:357. 1989, 

head one men ele To ae Bot. Zhurn. 75:225. 1990. 

. Riviére, Bull. Soc. Acclim. sér. 3, hae 1878. 

Drenavactas hyum a atum (Nees) Keng fC var. ¢ ramble, Ann. Bot. Gard. Calcutta 7:13. 1896; emend. 
Pandey, D.N. Tewari, Monogr. Bamboo 84. | 1903 TYPE: INDIA ul FAR PRADESH: Jaunsar Bawa, 1879, Bagshawe 
s.n. (LECTOTYPE designated here, K, superseding previous citation of type by Chao & Renvoize, Kew Bull. 
44:358. 1989, 


Further Sensseiey INDIA. ae Pradesh: Jaunsar, Aug 1878, Bagshawe 6608(K): Jaunsar, Comm. Brandis s.n. a 
Ss) u K). 


(flowers) only, t 


Drepanostachyum falcatum var. sengteeanum and other similar bamboos 

Having excluded the name Drepanostachyum falcatum var. glomeratum from the 
plants once cultivated at Kew and currently in horticultural use in the USA, it is neces- 
sary to consider other possible names. 

Arundinaria interrupta Trin. from Nepal has variable development of light, white 
hairs on the culm nodes, noted in the description. Culm sheaths are not present in the 


—_— 


STAPLETON CNCTECCANIINA ogee 


type, but recent collections from Nepal from a similar location have, in addition to simi- 
lar white hairs on the culm nodes, the distinctive sparse white hairs on most of the culm 
sheath and the mainly glabrous culm sheath bases of other collections of D. falcatum. 
Even the lectotype of A. falcata has some white hairs on the persistent culm sheath base, 
and this character was also given in Tewari (1993) for the type variety. The possibility of 
Arundinaria interrupta Trin. representing a separate species was raised (Stapleton 1994), 
but the new collections suggest that it should remain a synonym of D. falcatum. 

Other bamboos with hairs at the base of the culm sheath have been collected else- 
where in the Himalayas. Himalayacalamus fimbriatus Stapleton also has culm sheaths 
with a basal ring of hairs and a fimbriate ligule, but the larger stature, the much larger 
leaves, as well as the asymmetry of the culm sheath with its broader ligule and less sca- 
brous interior distinguish it clearly. Drepanostachyum annulatum Stapleton from Bhutan 
also has a ring of hairs at the base of the culm sheath, but the hairs are darker, longer, 
more upright and ina narrower ring. That species also differs in having dark brown culm 
sheath hairs, especially on the distal third of the culm sheath, which also bears erect, 
basally scabrous oral setae when young, as well as darker brown, longer cilia of up to 
1.5mm on the edges. It also has well-developed leaf sheath auricles, and thicker culm wax. 
In addition the ligules are neither as delicate nor as laciniate, and the culm sheath is less 
pubescent on the inside at the base of the ligule and below it. Although it shares the pos- 
session of a basal ring of culm sheath hairs with H. fimbriatus and D. annulatum, the 
bamboo currently cultivated in the US, recently described as D. falcatum var. sengteeanum 
(Stapleton 2006) is clearly much closer to the type of D.falcatum. 

The 2-3 flowered florets and the densely pubescent culm sheath interior apex of var. 
sengteeanum are sufficient to place this cultivated bamboo indisputably in 
Drepanostachyum. The absence of oral setae and the distribution of hairs on the culm 
sheaths are sufficient to separate it clearly from all previously described taxa. Following 
current taxonomic concepts in bamboos, culm sheath pubescence and presence of oral 
setae are important and consistent specific characters. They are usually supported by 
minor differences in spikelet or floret detail, while leaf sheath and blade characters are 
inherently more variable. However, in Drepanostachyum more variety is seen within spe- 
cies than in most other bamboo genera, and it was felt inappropriate to describe this taxon 
as a new species, especially as its origin remains unknown. The florets and spikelets of D. 
falcatum var. sengteeanumare more scabrous than those of the type variety and the palea 
is less bifid. The rhachilla segments are slightly longer and have rather different apical 
pubescence: the beard is mainly restricted to the distal 0.25mm of the rhachilla, while in 
the type of D. falcatum the distal 1mm becomes progressively more puberulent. The 
lemma and palea are more densely scabrous than any other material of D.falcatum from 
the Western Himalayas, and approach the state seen in other species from the Eastern 
Himalayas, suggesting a Nepalese or Sikkimese origin. 


ACKNOWLEDGMENTS 


The author would like to express his deep gratitude to Seng-Tee Lee, in whose honor the 
variety D.falcatum var. sengteeanum was named, for funding the Sino- Himalayan Bamboo 
HSO post at Kew for six years from 1994 to 2000. Staff at LIV are thanked for loan of Royle 
specimens. Gib Cooper is thanked for initially sending material from Oregon. The Ameri- 
can Bamboo Society is thanked for supporting travel to lecture and collect cultivated mate- 
rial in California. Reviewers are thanked for helpful comments on the manuscript. 


1086 BRIT.ORG/SIDA 22(2 


REFERENCES 


Bean, W.J. 1907. The flowering of cultivated bamboos. Bull. Misc. Inform. Kew 1907:228-233. 

Cao, CS. and S.A. Renvoize. 1989. A revision of the species described under Arundinaria (Gramineae) 
in southeast Asia and Africa. Kew Bull. 44:349-367, 

Game, J.S. 1896. The Bambuseae of British India. Ann. Roy. Bot. Gard. (Calcutta) 7:1—133. 

Munro, W. 1868.A monograph of the Bambusaceae. Trans. Linn. Soc. London 26:1-157. 

NaiTHANI, H.B. and S. CHanpra. 1998. Flowering of a bamboo (Drepanostachyum falcatum). Indian For- 
ester 124:663-666. 

Nees Ab Esenseck, C.G, 1834. Bambuseae Brasilienses. Linnaea 9:461-494, 

Neal, S.S. and H.B. NarrHani. 1994, Handbook of Indian bamboos. Oriental Enterprises, Dehra Dun, India. 

Rovie, J.F. 1839. Illustrations of the botany of the Himalayan Mountains. Allen, London 

SEETHALAKSHMI, K.K. and M.S. MuktesH Kumar. 1998. Bamboos of India: A compendium. Kerala Forest Re- 
search Institute & INBAR. 

Star, O. 1904. Himalayan bamboos. Gardeners’ Chronicle May 14:305-306; May 21:325-326; June 
4:356. 

STAPLETON, C.M.A.1994.The bamboos of Nepal and cs Part Ill: Drepanostachyum, Himalayacalamus, 

(Gramineae: Poaceae, Bambusoideae). 


} 


Ampelocalamus, Neomicrocalamus, and Chi 
Edinburgh J. Bot. 51:301-330 

STAPLETON, C.M.A. 2006. New taxa and combinations in cultivated bamboos (Poaceae-Bambusoideae). 
Sida 22:331-332. 

Tewari, D.N. 1993.A monograph on bamboo. International Book Distributors, Dehra Dun, India. 


REJECTION OF LECTOIYEIRIGATION OF 
ASTER AMELLUS (ASTERACEAE: ASTEREAE) AND 
SELECTION OF A NEW LECTOLYPE 


John C.Semple 


Department of Biology 
University of Waterloo 

00, Ontario, CANADA N2L 3G] 
jcsemple@sciborg.uwaterloo.ca 


Water, 


ABSTRACT 


Selection of 997.18 (LINN) as the lectotype of Aster amellus L. is rejected because the specimen is in conflict with 


the peat aes A new Oy, pe is designated f rom 1 authentic sian seen by Linnaeus. oe the original 
or conservation or redefining the 


eee of the genus Aster L. Herb, is inn. 997, 16 (1 -INN) is designated the new lectotype o Aster cls 


RESUMEN 


Se rechaza la seleccién del pliego 997.18 (LINN) como lectotipo de Aster amellus L. porque el espécimen esta en 
conflicto con i Protloeo, Se eclen un nuevo lectotipo a pare de malete auténtico visto ES Linnaeus. oo 
rechazo de la ipi la seleccion de vacion 
o redefinicion del cone del ae Aster L. El Biesollieit Linn, 907. 16 (LINN) se eee nuevo lectotipo de 
Aster amellus. 


Semple (1998) designated Herb. Linn. 997.18 (LINN) as the lectotype of Aster amellus L. 
(Astereaceae: Astereae). Recently, this lectotypification was found to be flawed and would 
result in a change in the application of the name Aster of which A. amellus is the 
generitype. The simplest solution to this situation is to reject the original lectotypification 
because the selection was in conflict with the protologue. An alternative solution would 
be to propose conservation of A. amellus with a conserved type, which does not seem 
necessary in this case. 

In choosing 997.18 (LINN) as the lectotype for Aster amellus for the Linnaean Typifi- 
cation Project (Semple 1998), I worked from a set of small photographs I had taken in 
1984 during visits to the Linnaean Herbarium (LINN) and the British Museum of Natu- 
ral History (BM) and from additional notes made in 1994 on numerous authentic speci- 
mens of asters and goldenrods. Other authentic specimens were rejected as possible types 
then, and additional specimens and photographs have subsequently come to my atten- 
tion. 997.18 (LINN) is here rejected as the lectotype for Aster amellus on the basis of being 
in conflict with the protologue. Article 9.17(b) of the ICBN (McNeill et al. 2006) permits 
the supersession of a lectotype, if it is in serious conflict with the protologue, and an- 
other element that is not in conflict is available. At least one specimen that does not con- 
flict with the protologue, Herb. Linn. 997.16 (LINN), is available. 

In the protologue of Aster amellus, Linnaeus (1753, p.873) indicated the occurrence 
of some variation in leaf and phyllary traits, and he noted the range as “Habitat in Europae 
australis asperis collibus,” that is, native to rugged hills in southern Europe. Leaves were 
described as “lanceolatis obtusis scabris trinervis integris” and phyllaries as “calycinis 
obtusis” in the diagnosis. LINN 997.18 has linear, uninervate rather than lanceolate 
trinervate leaves and linear acute rather than obtuse phyllaries. The small black and white 


SIDA 22(2): 1087— 1089. 2006 


1088 BRIT.ORG/SIDA 22(2) 


photograph of 997.18 (LINN) that | worked from was cropped and lacked critical annota- 
tions that are obvious in the colored image of the specimen currently posted on the Lin- 
naean Typification Project web site http://interntnhm.ac.uk/jdsml/research-curation/ 
projects/linnaean-typification /index.dsml. Upon seeing this image, | knew immediately 
that |had made a mistake in my choice of lectotype for Aster amellus. The specimen was 
annotated “non A. Amellus” by James Edward Smith (Charlie Jarvis, pers. comm.), who 
bought Linnaeus’ herbarium and moved it to England, and as “Calimeris A.G” by Asa 
Gray. Clearly, these annotations are critical because they indicate the practical conse- 
quences of retaining 997.18 (LINN) as the type of Aster amellus, the generitype; the mod- 
ern strict application of the name Aster (see Nesom 1994; Semple et al. 2002) would fun- 
damentally have to be changed. 997.18 (LINN) is likely a member the Galatella Cass. / 
Crinitaria Cass. complex rather than Aster 

The second paragraph of the protologue deals with the first synonym Linnaeus cited 
from Hortus Cliffortianus. The voucher of the synonym cited as “Hort. Cliff. 407.” is Clif- 
ford Herb. No. 407 (BM!) and was diagnosed by Linnaeus as having leaves “lanceolatis 
scabris semiamplexicaulisbus serratus” and phyllaries as “lanceolatis.” Linnaeus noted 
that the “Hort. Cliff.” reference was previously cited in “Royen” and “Gmelin.” Clifford 
Herb. No. 407 (BM) is a possible choice for lectotype for Aster amellus. The specimen is a 
fragment, and in leaf features it is similar to Herb. Linn. 997.17 (LINN), a specimen of A. 
pyrenaeus Desf. ex DC. in Lam. & DC. The Clifford herbarium specimen is rejected as a 
choice for the new lectotype of A.amellus. 


—y 


Retypification 

Herb. Linn. 997.16 (LINN) is also an authentic specimen and is annotated “5 Amellus” 
likely by Linnaeus himself (Charlie Jarvis, pers. comm.). | have no data indicating I saw 
this specimen in 1984 or 1994. A photograph of this specimen was given to me after 1998 
by Paul Van Faasen, who took the picture. The plant illustrated is very similar to plants 
in my research collection of Aster amellus collected for me in Switzerland by Dr. Erich 
Weber. 997.16 (LINN) and the Weber collections match the description of Aster amellus 
in the diagnosis of the protologue in both leaf and phyllary traits. These correspond well 
with the description and figures of A. amellus in Wagenitz (1979) and the description of 
the species in Merxmiuller, Schreiber and Yeo (1976). Therefore, Herb. Linn. 997.16 (LINN) 
is here designated the new lectotype of Aster amellus. 


Aster amellus L., Sp. Pl. 2:873. 1753. Type: EUROPE: habitat in Europae australis asperis co oe [native 
to rugged hills in southern Europel, Herb. Linn. 997.16 (LECTOTYPE here designated as new: LINN 


ACKNOWLEDGMENTS 


This work was supported by a Natural Sciences and Engineering Research Council of 
Canada Discovery Grant to the author. Charlie Jarvis of the British Museum of Natural 
History is gratefully acknowledged for his assistance with Linnaean collections at BM 
and LINN. Fred R. Barrie and Charlie Jarvis are thanked for their useful and timely re- 
views of the submitted manuscript. lam grateful to Paul Van Faasen for having chosen to 
give me his extensive collection of photographs of type species of asters and copies of 
species protologues. 


SEMPLE, RE-LECTOTYPIFICATION OF ASTER AMELLUS 1089 


REFERENCES 


Linnaeus, C.1753. Species Plantarum. Stockholm. 

McNett, J., FR. Barrie, HM. Buroet, V. DEmMouLIN, D.L. HAwKSworTH, K. MARHOLD, D.H. Nicotson, J. PRADO, PC. Sitva, 
J.E. Skoa, J.H. Wiersema, and NJ. Turcanp. (eds.) 2006. International Code of Botanical Nomenclature 
(Vienna Code)...Adopted by the Seventeenth International Botanical Congress Vienna Austria, 
July 2005. (Regnum Veg. 146). Koeltz Scientific Books, KOnigstein, Germany 

Merxmutter, H., A. ScHreiber and PF. Yeo. 1976. Aster. In: Tutin, T.G.,V.H. Heywood, N.A. Burges, D.M. Moore, 
D.H.Valentine, S.M.Walters and D.A.Webb, eds.Flora Europaea.Vol.4. Plantaginaceae to Compositae 
(and Rubiaceae). Cambridge University Press, Cambridge, U.K. 

Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the 
New World species. Phytologia 77:141-297. 

Sempte, J.C. 1998. Typification of Aster amellus, A.dumosus, A.novae-angliae, A. puniceus, A. sibiricus, Sol- 
idago flexicaulis, S. lanceolata and S. latifolia. |n: Jarvis, C.E. and N.Turland, eds. Typification of Lin- 
naean names in the Asteraceae (Compositae). Taxon 47:347-370. 

Semee, J.C., S.B. Heard and L. Brouiter. 2002. Cultivated and native asters of Ontario (Compositae: 
Astereae):Aster L. (including Asteromoea Blume, Diplactis Raf.and Kalimeris (Cass.) Cass.), Callistephus 
Cass., Galatella Cass., Doellingeria Nees, Oclemena E.L. Greene, Eurybia (Cass.) S.F. Gray, Canadanthus 
Nesom, and Symphyotrichum Nees (including Virgulus Raf.). Univ. Waterloo Biol. Ser. 41:1-134. 

Wacenitz, G. 1979. Aster. In: Hegi, G., ed. Illustrierte Flora von Mittel-Europa. Band 6(4), Compositae. 
1:35-71. Hehman Verlag, Munich. 


1090 BRIT.ORG/SIDA 22(2) 


Book NOTICE 


S. InpbeRIt (ed). 2005. Invasive Plants: Ecological and Agricultural Aspects. (ISBN 3-7643- 
7137-4; 978-3-7643-7137-1, hbk.). Birkhauser Verlag, PO. Box 133, CH-4010 Basel, 
SWITZERLAND. (Orders: Springer, 333 Meadowlands Parkway, Secaucus, NJ 07094, 
U.S.A.; 800-SPRINGER:; fax 201-348-4505: WW w.spt inger.com). $159.00, 283 pp., 
illustrations, 6 1/2" x 91/4". 

A set of studies toward understanding the biology of invasive species, comprising 17 papers written by an inter- 


national group of invasions biologists. The chapter titles give a good view of the content. 


* In search of an operational lexicon for biological invasions. 

* The ecology of biological invasions: past, present, and future. 

* Ecological niche models and the geography of biological invasions: a review and a novel applica- 
tion. 

* Importance of species replication in understanding plant invasions into North American grass- 
lands 

* Residence time determines the distribution of alien plants 

* The relationship between community diversity and exotic plants: cause or consequence of inva- 
sion? 

* Invasive plants: the process within wetland ecosystems. 

* Understanding invasions:the rise and fall of diffuse knapweed (Centaurea diffusa) in North America. 

* Defining a regional approach for invasive plant research and management. 

* Allelopathy as a mechanism for resisting invasion: the case of Polygonella myriophylla. 

* Ecology and management of an exotic weed Phalaris minor. 

* Reducing agroecosystem vulnerability to weed invasion. 

* Tillage intensity affects weed communities in agroecosystems. 

* Distribution and biological control of Chromolaena odorata. 

* Biology and anthropology of plant invasions. 

* Characters, significance, and human dimension of global invasive weeds. 

* Environmental consequences and economic costs of alien species. 


—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, UX 76102-4068, U.S.A. 


SIDA 22(2): 1090. 2006 


TRICHOME MORPHOLOGY IN SELECTED MEXICAN 
RED OAK SPECIES (QUERCUS SECTION LOBATAE) 


M. Lucia Vazquez! 
Department of Plant Biology 
Cornell University 
Ithaca, New York 14850, U.S.A. 
vazquez.lucia@uis.edu 


ABSTRACT 
Mature leaves and twigs noe oy red oak Species, meaty nas to oes were examinee with SEM to 
assess trichome eee ee i g g glandular. Described 
for the first ti fi icul df icul d d ltipl 1] trichomes. 
Furthermore, tl ick i I 1 for the fi ime i 1 oal llate, fused stellate and glandular 
branched. Tl | 1 f l | | lti li vhile 


f. ery | cf ese eee ea ay see aoe 4 1p; 


the most frequent types on the eee llea 
trichomes. In contrast to the significant anes ees variation found in oak leaf a ‘cichomle ee 
within each species are Eeacanye invaria ble ane t De useful in the iain canes of most species. In summary, 
together with veg i gy | ional information for 


-_ 


the identification and chactenause of Mexican oak sp 


RESUMEN 


Hojas maduras y ramillas de treinta especies de encinos rojos, principalmente endémicos a Mexico, fueron 
mas con pena ae Secu onica de BarEdo pana evaluar la variacion morfolégica de tricomas. En este 
glandulares. pe estos, se ‘ii iben por 


tr abajo of 


primera vez para encinos jesus tricomas s fasciculadoc contorto, lagen ado Gsiado y estr \demas 
se reportan por panes para j p llado fusi y glandu- 
larramifieado.| r ic] j ill | fasciculado sésil y el multiradiado, mientras 
S| ape | al : ] . ] : Ane es ‘ leael Beil £. G la- tin} ] 

f ic : r y el 

fies A ] ‘ fologia fol 
L i=) Oo .o L 

en cada especie son te por | utiles en la identificacion de la mayoria de las especies. En 


resumen, los caracteres vegetativos y reproductivos junto con la morfologia de trichomes proporcionan 
informacion valiosa para la identificacion y caracterizacién de las especies Mexicanas de encino. 


INTRODUCTION 


The genus Quercus (Fagaceae) consists of approximately +00 species of trees and shrubs 
distributed in temperate and subtropical regions of the Northern Hemisphere. In the 
Americas, Quercus is found from southern Canada to Colombia with representatives of 
three sections within the subgenus Quercus: Quercus (white oaks), Protobalanus (inter- 
mediate oaks), and Lobatae (red oaks) (Nixon 1993). Due to the large diversity of white 
and red oaks, Mexico is considered the center of diversity for the genus in the New World 
(Rzedowski 1965; Nixon 1993). 

he number of red oak species has been estimated around 195 (Jensen 1997) with 
about 55 species endemic to Mexico (Nixon 1993). In his treatment on the American oaks, 
Trelease (1924) segregated the red oak species into 73 series and also proposed a classifi- 
cation based on leaf and fruit morphology. However, his hypothesis on the relationships 
of red oak species has not been tested with phylogenetic methods. Although the mono- 


‘Current address: Biology Department, University of Illinois Springfield at Springfield, Illinois, 62794, U.S.A. 


SIDA 22(2): 1091- 1110. 2006 


1092 BRIT.ORG/SIDA 22(2) 


phyly of the red oak section is well supported (Nixon 1984; Manos 1999), an intrasectional 
phylogeny for the red oaks is yet to be conducted. As in other oak groups, Mexican oak 
species are difficult to study because most of them show significant amounts of foliar 
polymorphism. Species identity becomes even more complex when either hybridization 
or introgression has taken place. Therefore, taxonomic problems in many oak species 
persist, particularly in those species that have similar or overlapping leaf morphology. 
The utility of trichome morphology for species identification has been demonstrated 
in several plant families such as Brassicaceae, Geraniaceae, and Sterculiaceae (Inamdar 
& Rao 1983; Metcalfe & Chalk 1950; Oosthuizen 1983; Shanmuka Rao 1987). In oaks, tri- 
chome characterization has been carried out for several eastern North American, Asian, 
and European species. The first study on trichome morphology of eastern North Ameri- 
can oak species was reported by Dyal in 1936, who classified them, based on secretory 
function, into glandular and nonglandular. Camus (1934-1953) described seven trichome 
types that fall into the glandular and nonglandular categories previously described by 
Dyal (1936). Hardin (1976) published the first comprehensive work on oak trichome mor- 
phology for 58 eastern United States (US) oak species and recognized 10 trichome types. 
Thomson and Mohlenbrock (1979), Jones (1986), and Manos (1993b) conducted similar 
studies on other U.S. oak species. Trichome morphology of European and Mediterranean 
oak species has also been extensively documented by several authors (Olsson 1976; 
Kissling 1977, 1993; Safou & Saint-Martin 1989; Gellini et al. 1992; Llamas et al. 1995; 
Bussotti & Grossoni 1997). Trichome morphology, together with other vegetative and re- 
productive characters, has been used to discriminate several USS. oak species, as well as a 
few Mexican species distributed in the northwest of the country (Tucker 1952; Tucker & 
Muller 1957; Nixon & Steele 1981; Spellenberg 1992, 1998; Nixon & Muller 1993; Bacon & 
Spellenberg 1996). However, to date, no in-depth research on the trichome morphology 
of Mexican taxa has been carried out. Several regional taxonomic treatments of Mexican 
oak species have been conducted (Gonzalez 1986; Valencia 1989; Vazquez 1992, 2000; 
Spellenberg et al. 1998; Spellenberg 2001), but none of them has characterized trichome 
diversity. Only a few authors have discussed the potential utility of trichome types for 
identification of Mexican species (Mc Vaugh 1974; Rzedowski & Rzedowski 1979; Vazquez 
1992). Accordingly, the objective of this study is to characterize the trichome morphol- 
ogy from twigs and leaves from thirty Mexican red oaks, mainly endemic to Mexico, us- 
ing scanning electron microscopy (SEM). Detailed examination of these species revealed 
the presence of ten trichome types: four of these trichomes have been previously described, 
three of them represent entirely new trichome types, and the remaining three are tri- 
chome types not previously documented for red oaks. Each species examined shows a 
particular trichome complement which, together with other vegetative and reproductive 
characters, provides useful information for species identification and characterization. 


MATERIALS AND METHODS 
Leaves and twigs from thirty species of red oak species chiefly endemic to Mexico were 
selected for examination of trichome morphology under SEM. Nixon (1993) has reported 
41 red oak species endemic to Mexico, of which, twenty-one have persistent indument 
and were sampled in this study. Two additional species with persistent indument (Q. 
castanea and Q crassifolia) whose range extends to Central America were included in 
this study because they are widely distributed in Mexico. For comparison, six glabrous 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 


Tape 1. Red oak species (Quercus) examined with SEM for trichome morphological variation. Voucher speci- 
mens are deposited at Bailey Hortorium, Cornell University (BH 


= 


Taxa 


Voucher specimen 


Q. acherdophylla Trelease 
Q. affinis Scheidweiler 

Q. aristata Hooker & Arnott 

Q. candicans Nee 

Q. castanea 

Q. coahuilensis Nixon & Muller 

Q. coccolobifolia Trelease 

Q. conzattii Trelease 

Q. crassifolia Humboldt & ly 


Q. crassipes Humboldt & Bon plan 
Q. depressa Humboldt & :Sonpbnd 
Q. auido ia von Seem 


lysopt rylla Ber itt lalit 
QO ee 
Q. fulva Liebmann 
Q.gentryi Muller 
Q. hintonii Warbur 


Vdzquez et al. 3065 

Muller 9716 

Martinez 31, Breedlove 1640 
Vdzquez et a 

Rzedowski 25361 

Stewart 924 

Vazquez & Phillips 3084 
Muller 9420 

Vazquez et al. 3049 
Vazquez et al. 3037, 3070 
Vdzquez & Tenorio 118 
Muller 3579 

Nixon s.n. (Oct. 1999) 

Nixon 4034, Vazquez et al. 3011 
Vazquez et al. 3097 

McVau eee 256 . 

Vazquez 


al.: 
cron ne 2336 
Vazquez et al. 3069, 3109 
Ven & Fetrqdgq 22322 


Q. hintoniorum Nixon & Muller 
Q. hirtifolia Vazquez, Valencia & Nixon 
Q. hypoleucoides Camus 


Q. hypoxantha Trelease Cowan et al. 5402 

Q. mevaughii Spellenberg Bacon et al. 5284 
mexicana Humboldt & Bonpland Rzedowski 6374 

Q. planipocula Trelease McVaugh12136 

Q. radiata Trelease Rzedowski 23018 

Q. salicifolia Né McVaugh 25456 

Q. scytophylla Liebman Vazquez et al. 3035 


Q. sideroxyla ree & Bonpland Vazquez et al. 3089, Bacon et al. 5304 
Q. tarahumara Spellenberg, Bacon & Breedlove Bacon et al. 5307, 5375 
Q. urbanii Trelease Vazquez et al. 3027 


red oak species endemic to Mexico were included. Samples of the materialexamined were 
gathered from field collections and from herbarium specimens deposited at the Bailey 
Hortorium, Cornell University (BH) (Table D. 

Twenty to thirty specimens per species were initially examined under dissecting and 
light microscopy to survey the intra- and interspecific variation in trichome morphol- 
ogy. A more detailed examination was carried out under SEM in representative 
subsamples. About one square cm of the abaxial surface of a dry leaf or one cm of twig 
was mounted on an aluminum stub with no prior treatment. Previous studies have shown 
no difference in the preservation of nonglandular trichome structure using either fresh 
or dry tissue (Hardin 1976). Specimens mounted on stubs were sputter coated with 30 
nm of gold palladium using a BAL-TEC sputter coater. Samples were observed under a 
Zeiss electron microscope, model LEO DSM 960, using working distances between 12 and 
18 mm, voltages of 3.0 or 4.0 kV, and apertures of 300 um or 400 um depending on the 
sample and magnification. Most trichome micrographs were taken at magnifications of 
200, although smaller trichomes required magnifications of 450-500. 


1094 BRIT.ORG/SIDA 22 


2) 


Tasve 2. Trichome types on twigs of Mexican red oak species (Quercus). 


Species Glandular Nonglandular 
Simple Branched Fasciculate Fasciculate Multiradiate Simple 
uniseriate sessile simple —_stipitate Stellate 

Q. acherdophylla x Xx 
Q. affinis X 
Q. aristata xX 
Q. candicans Xx Xx x 
Q. castanea Xx 
Q. coahuilensis x x 
Q. coccolobitolia xX x 
Q. conzattii x X 
Q. crassifolia xX Xx x 
Q. crassipes X X 
Q. depressa X 
Q. durifolia xX Xx 
Q. x dysophylla X Xx 
Q. eduardii x Xx 
Q. fulva x X 

). gentryli Xx 
Q. hintonii X X 
Q. hintoniorum X X 
Q. hirtifolia Xx x 
Q. hypoleucoides Xx x 
Q. hypoxantha X xX 
Q. mevaughil Xx Xx 
Q. mexicana X Xx 
Q. planipocula Xx 
Q. radiata X 
Q. salicifolia x 
Q. scytophylla Xx x 
Q. sideroxyla Xx Xx 
Q. tarahumara X 
Q. urbanii X X 

RESULTS 


Trichome nomenclature follows Hardin (1976), where each cell composing a nonglandular 
trichome is called a “ray” and trichome types are named based on the degree of ray fu- 
sion, ray orientation, and overall morphology. However, additional terms have been in- 
troduced to describe trichome types not previously reported in the literature. 

Two general trichome categories are used to indicate secretory functions and nature 
of the cell wall: 1) glandular trichomes have a glistening appearance and thin cell walls 
that collapse upon drying, and 2) nonglandular trichomes lack a glistening appearance 
and possess thick cell walls. Examination of twigs and abaxial leaf surfaces of thirty red 
oak species resulted in two types of glandular trichomes and eight nonglandular trichome 
types (Tables 2 and 3). 

Glandular trichomes 

1. Simple uniseriate (Fig. 1A, 1B).—This type consists of 2-7 united cells with slight 

constrictions along the trichome length. This trichome type was commonly found on 


—_— 


Taace 3. Trichome types in oak species (Quercus) with a persistent indumentum on the abaxial leaf surfaces. 


Species Glandular Nonglandular 
Simple Branched Fasciculate Fasciculate Fasciculate Fasciculate Multi- Simple Fused Multiple 
uniseriate sessile sessile sessile stipitate radiate stellate stellate stellate 
simple contorted crested 

Q. acherdophylla XV 
Q. affinis Xv 
Q. aristata X Xx 
Q. candicans Xx x Xx X 
Q.castan X Xx 
Q. coahuilensis xX x 
Q. coccolobifolia x Xv 
Q. conzattii X 
Q. crassifolia x x X 
Q. crassipes X x 
Q. depress Xv 
Q. durifolia X x 

. x dysophylla x x 
Q. eduardii xX 
Q. fulva x x 
Q.gentryi Xx X Xs 
Q. hintonii x x x 
Q. hintoniorum xX Xv x 
Q. hirtifolia x xX 
Q.hypoleucoides =X xX 
Q. hypoxantha x x 
Q. mevaughii X X 
Q. mexicana xX X 
Q. planipocula x x 
Q. radiata xX x 
Q. salicifolia XS XS 
Q.scytophylla x X 
Q. sideroxyla x x X 

tarahumara xX x X 
Q. urbanii xX X 


s=scattered 


v=trichomes on secondary veins 


SNIYINO NI ADOTOHdYOW IWOHDIIYL ‘ZINDZVA 


S60L 


1096 BRIT.ORG/SIDA 22(2) 


\ 


Fic. 1.A. Thi | Wi | 1] +t tn lobifolia. x 40.B. Glandul . 1 


leaf surface of Q. hintoniorum, x 500. C. Glandular t hed trict | surf f Q. salicifolia, x 500. D. A variation of 
the glandular I hed trict the abaxial | f Q. salicifolia, x 500. 


the abaxial leaf surfaces of twenty-one species, forming a sparse and deciduous 
~matu- 


indumentum. In most species examined, trichome density decreased during lea 
ration and senescence; however, Quercus coccolobifolia showed persistent simple uniseriate 
trichomes on the abaxial leaf surface. This trichome type was found on the twigs of only 
three species: Q. acherdophylla, Q. coccolobifolia, and Q. crassifolia (Table 2). 

2. Branched (Fig. 1C, lD).—This type consists of 2-3 cells united in either a V or a Y 
shape; however, under the dissecting microscope this trichome type looks like two uni- 
cellular simple trichomes lying next to each other. This trichome type was found exclu- 


sively on abaxial surfaces of Q. salicifolia leaves. 
Nonglandular trichomes 
Fasciculate sessile trichomes.—These 


e consist of several rays fused at the point of attach- 
ment to the epidermis. The rays are either short and completely erect (Fig. 2A), or long 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1097 


rc . nl eee oe 
Fic. 2. A t trichome on 


a. 4 | th |} oe | | 
chome on the abaxial leaf surface of Q. crassifolia, < 200. C. F 
mexicana, 200 iculat il torted trict the abaxial leaf surf. f Q. eduardii < 200. 


f Quercus aristata, X 200. B. Fasciculate sessile simple tri- 
a 1 1 th L a i | £ £. [ph 
Q. 


and tortuous diverging in different directions (Fig. 2B). Three types of fasciculate sessile 
trichomes are recognized in the species examined. 

1. Fasciculate sessile simple (Fig. 2A, 2B).—The fasciculate sessile trichome described 
by Hardin (1976; 1979a) and Jones (1986) is termed here fasciculate sessile simple to dis- 
tinguish it from the two additional subtypes listed below. This trichome type was found 
on twigs and/or on the abaxial leaf surfaces of several species examined. It occurs on 
twigs of the following species: Q. candicans, Q. castanea, Q. coahuilensis, Q. coccolobifolia, 
QO. conzattii, Q. crassifolia, Q. crassipes, Q. durifolia, Q. x dysophylla, Q. eduardii, Q. fulva, Q. 
hintonii, Q. hintoniorum, Q. hirtifolia, Q. hypoleucoides, Q. hypoxantha, Q. mcvaughii, Q. 
mexicana, Q. planipocula, Q. radiata, Q. scytophylla, Q. sideroxyla, Q. tarahumara and Q. 
urbanii. Species bearing this trichome type on abaxial leaf surfaces are Q. aristata, Q. 
castanea, Q. coahuilensis, Q. crassifolia, Q. hintonii, Q. hirtifolia, Q. hypoxantha, Q. 
mcvaughii, Q. radiata, and Q. tarahumara (Table 2). 


1098 BRIT.ORG/SIDA 22(2) 


2. Fasciculate sessile contorted (Figs. 2C, 2D).—This study describes this trichome 
type for the first time. Contorted trichomes are similar to the fasciculate sessile simple 
trichome described above except that the rays curl down and then twist around the tri- 
chome axis. This ray arrangement gives the indumentum a punctate appearance when 
observed with the naked eye or at low magnification. This trichome type has been found 
on the abaxial leaf surface of only two species of Mexican red oaks: Q. mexicana and Q. 
eduardii. 

3. Fasciculate sessile crested (Fig. 3A).—This is a new trichome type that consists of 
rays united along a longitudinal axis oriented parallel to the epidermis. The united rays 
result in a structure that resembles a crest. Trichomes of this type were found only on the 
abaxial surface of some Q. sideroxyla specimens. 

4. Fasciculate stipitate (Figs. 3B, 3C).—Morphologically, this trichome type is similar 
to the fasciculate sessile simple except that the basal parts of the ae fuse, forming a 
stipe. The degree of ray fusion varies among species from 1/12 to 1/4 the total ray length. 
Ray length and degree of divergence are also different; some species display short and 
straight rays (Fig. 3B) while others show long curly rays that become intertwined with 
adjacent rays (Fig. 3C). Ray number is variable within and among species, although in- 
traspecific ranges are low. Fasciculate stipitate trichomes form a persistent indumentum 
on the abaxial leaf surface of the following species: Q. conzattii, Q. crassifolia, Q. crassipes, 
Q. x dysophylla, Q. hintonii, Q. hirtifolia, Q. planipocula, Q. tarahumara,and Q. urbanii. In 
Q. aristata, fasciculate stipitate trichomes are deciduous and scattered on the abaxial leaf 
surface. In many otherwise glabrous species, such as Q. acherdophylla, Q. affinis, Q. 
coccolobifolia, Q. depressa, and Q. hintoniorum, only a few fasciculate stipitate trichomes 
remain attached to the epidermis at the secondary vein axils. Fasciculate stipitate tri- 
chomes on twigs were less common and found only in Q. aristata, Q. coahuilensis, Q. 
crassifolia, Q. x dysophylla and Q. hirtifolia. 

5. Multiradiate (Figs. 3D, 4A, 4B).—This trichome type is characterized by the diver- 
gence of rays in different directions from the central axis. In the species examined, the 
rays differ not only in length but also in thickness. Multiradiate trichomes were found on 
both twigs and abaxial leaf surfaces of Q. candicans, Q. durifolia, Q. fulva,Q. hintoniorum, 
Q. hypoleucoides, Q. scytophylla, and Q. a This trichome type was found only on 

the twigs of the following species: Q. affinis, Q. conzattii, Q. crassipes, Q. depressa, Q. 
eduardii,Q. gentryi, Q. hintonii,Q. hypoxantha, Q. mcvaughii, Q. mexicana, and Q. urbanii. 
Multiradiate trichomes occurring on abaxial leaf surfaces usually form a persistent 
indument, however, in Q. salicifolia these trichomes are found scattered sparsely across 
the lamina. 

6. Simple stellate (Figs. 4C, 4D, 5A).—This trichome type consists of rays appressed 
to the lamina, usually fused at their base, forming a structure that resembles a starfish. 
Simple stellate trichomes are composed of seven to fifteen rays, which vary in length, 
thickness, and size. This trichome type was previously described by Hardin (1976) and 
Jones (1986) under the term stellate. Here the term simple stellate is used to set it apart 
from the multiple stellate and fused stellate also found in this study. Simple stellate tri- 
chomes were found on the twigs of Q. acherdophylla, and Q. candicans, as well as on the 
abaxial leaf surfaces of Q. candicans, Q. durifolia and Q. gentryi. 

7. Multiple stellate (Fig. 5B).—This new trichome type consists of 2-3 stellate tri- 
chomes joined together forming a trichome cluster. It was found only on one species, scat- 
tered on the abaxial leaf surface of Q. gentryi. 


— 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1099 


Fic. 3. A. Fasciculat il tedt ial leaf surface of Q id la, * 450.B. Fasciculate stipitate trichome 
on nie abaxialleatutiace sche Ce aie C. Fasci ipitate trichome on ial leaf surface of Q. j 0.D. 
f Q. salicifolia, x 500. 


8. Fused stellate (Fig. 5C).—This trichome type previously described by Hardin (1979a) 
and Jones (1986) consis a of rays fused about one sixth of the ray length. This trichome 
type, not previously reported for red oaks, was found exclusively on the abaxial leaf sur- 
face of O. candicans. 

DISCUSSION 
The study of trichomes with SEM revealed significant differences in trichomes that ap- 
peared similar when viewed with dissecting microscopy. These differences have allowed 
a more accurate characterization of trichome types and have also led to the discovery of 
three new nonglandular trichome types: fasciculate contorted, fasciculate crested, and 
multiple stellate. Fasciculate contorted trichomes, although recognized asa different type 
by several authors (Mc Vaugh 1974; Gonzalez 1986), had been included under the stellate 


1100 BRIT.ORG/SIDA 22(2) 


Fic. 4.A. Multiradi oa ne eee peer Ree Jurifolia, x 300. B. Mul ej as eisai 
of Q. scytophylla, x 500. C. Simple stellate trich the adaxial leaf surf f Q. eduardii < 500. D. Simple stellate trichomes on 


the twigs of Q. acherdophylla 800. 


type. The second new trichome type, fasciculate simple crested, is found only on the 
abaxial leaf surface of some Q. sideroxyla specimens. The uncommon distribution of fas- 
ciculate crested trichomes suggests that they could represent a variation of the 
multiradiate type, which is an abundant trichome type in the indumentum of Q. 
sideroxyla. The third new trichome type, multiple stellate, is found ina scattered pattern 
exclusively on the abaxial leaf surface of Q.gentryi. Under the dissecting microscope this 
trichome resembles a multiradiate type but close examination with SEM reveals that it 
consists of two to three stellate trichomes fused together (Fig. 5B). Therefore, trichomes 
that appear multiradiate must be examined carefully to avoid inclusion of multiple stel- 
late types into this category. For example, Hardin (1979b) described a trichome type asa 
dense multiradiate, however, his corresponding illustration (Fig. 30) shows a morpho- 
logical resemblance to the multiple stellate type described here. 

In addition to the new trichome types found in this study, three types represent new 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1101 


Fic. 5. A. Simple stell ict the abaxial leaf surface of Q jurifolia X 250. B. Multiple stellate trichomes on lower leaf 
surface of Q. gentryii_450.C. Fused stell ict he abaxial leaf side of Q. candicans, < 200.D. Fasciculate sessile trichomes 


lleafetd f 0. coahuilensis. 150. 


reports for red oaks: stellate, fused stellate, and glandular branched. Previous research 
had indicated that stellate and fused stellate types occurred only in white oaks (Hardin 
1979a); however, this study found these trichome types in Q. candicans. Jones (1986) stated 
that stellate trichomes are “usually restricted to large veins or sometimes only the peti- 
ole.” In this research, stellate trichomes were found forming a dense and persistent 
indumentum on the abaxial leaf surfaces of the aforementioned species. The glandular 
branched type was rare and only found on the abaxial leaf surface of Q. salicifolia. This 
result supports Hardin's statement (1976) that this trichome type is not very common in 
red and white oaks. 

SEM was also useful in the accurate identification of previously known trichome 
types. For example, at low magnifications, trichomes on the abaxial leaf surface of Q. 
coahuilensis appear fasciculate stipitate, but examination under SEM has revealed that 
they are fasciculate sessile: the stipe-like structure is formed by the twisting of rays near 


1102 BRIT.ORG/SIDA 22(2) 


the trichome base (Fig. 5D). Similarly, detailed study of the trichome type denoted as 
“starred” in several descriptions of Mexican oak species (Bello & Labat 1987; Valencia 
1989; Vazquez 1992, 2000; Romero 1993) showed that these categories correspond to four 
different trichome types: fasciculat ile, fasciculate stipitate, fasciculate contorted, or 
multiradiate trichomes. Acknowledging the trichome variation included under the term 
“starred” will facilitate future characterization of Mexican oak taxa. 

Study of twigs under SEM has also revealed remarkable characteristics regarding 
the nature of the indumentum. Several taxonomic treatments of Mexican oak taxa 
(Gonzalez 1986; Valencia 1989; Vazquez 1992) imply that the twig indumentum consists 
of only one nonglandular trichome type. However, the results of this study indicate that 
the indumentum of nineteen of the species examined is composed of two to three tri- 
chome types arranged in overlapping layers (Fig. 6A). Of the thirty species examined, 
only nine have a twig indumentum composed of one nonglandular trichome type (Figs. 
OB, 6C). 

In general, species bearing more than one nonglandular trichome type on twigs and/ 
or leaves show several combinations of glandular and nonglandular trichomes, termed 
trichome complements. Glandular trichome types are not included in the trichome 
complements because, if present, they usually are deciduous. 


Trichome complements on twigs 

The most common trichome complement on twigs is multiradiate/fasciculate sessile 
simple and occurs in the following species: Q. conzattii, Q. crassipes, Q. durifolia, Q 
eduardii, Q. fulva, Q. hintoniti, Q. hinloniorum, Q. hypoleucoides, Q. hypoxantha, Q. 
mevaughii, Q. mexicana, Q. scytophylla, Q. sideroxyla, and Q. urbanii. Twigs with a tri- 
chome complement consisting of fasciculate sessile simple/fasciculate stipitate trichomes 
are found in Q. codhuilensis, fa) crassifolia, Q. x Cane and Q. hirtifolia. A species- 
specific trichome complement was found on twigs of Q. candicans (fasciculate sessile 
simple/multiradiate/simple stellate) (Table 2). 


Trichome complements on the abaxial surface of leaves 
The distribution of glandular trichomes on leaves showed different patterns. On one hand, 
the simple branched glandular trichome type is found exclusively on the abaxial leaf 
surface of Q. sdlicifolia in a scattered fashion (Fig. 6D). On the other hand, the simple 
uniseriate type is found on the abaxial leaf surface of most of the species examined. Gen- 
erally, this trichome is deciduous and usually not found on mature leaves: however, in Q. 
radiata and Q. coccolobifolia, these glandular trichomes are persistent in the form of 
mucilaginous droplets, which according to Uphof (1962) are the result of decaying glan- 
dular ae enn The common presence of simple uniseriate glandular trichomes has 
also been documented for many US and European oak species (Hardin 1976, 1979a; Lla- 
mas et al. 1995), and has also been found in the red oak species examined here. 
Trichome complements on the abaxial leaf surface are composed of two to four tri- 
chome types (Table 3). The trichome complement fasciculate sessile simple/fasciculate 
stipitate is found in the species Q. aristata, Q. crassifolia, Q. hintonii, Q. hirtifolia,and Q. tara 
pelcake aes epee tr ae eee oy en on the abaxial leaf surface 
), Q.durifolia (multiradiate/simple 
a ate), QO. hintoniorum (fasciculate stipitate/multiradiate), and Q. sideroxyla (fascicu- 
late sessile crested/multiradiate), (Table 3). Although the occurrence of more than one 
nonglandular trichome type on the abaxial surface has been documented for European 


—" 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1103 


6.6 A r . 1 yi Il L L 4 tn FY hd 200 R Cc = i: | = | l1nf4+\ L 1 
tate (right) trichomes on pa: at en 200.C. Multiradi ict the twigs of Q. affinis < 200.D. Scattered branched 
glandular trict f Q. salicifolia, < 200. 


(Safou & Saint-Martin 1989) and North American (Thomson & Mohlenbrock 1979) oak 
species, it has only been described for a few Mexican taxa. For example, Nixon and Muller 
(1993) described the unique trichome complement found in Q. hintoniorum and related 
species. 

The abaxial leaf surface of six of the included species is essentially glabrous except 
for clusters of fasciculate stipitate trichomes in the vein axils, or scattered multiradiate 
or stellate trichomes distributed throughout the lamina. Species with fasciculate stipi- 
tate trichomes restricted to vein axils are: Q. acherdophylla, Q. affinis, Q. depressa, and Q. 
coccolobifolia, although the later species also has abundant simple uniseriate trichomes 
throughout the abaxial surface. Scattered multiple stellate and multiradiate trichomes 
have been found only in Q. gentryi while dispersed multiradiate and branched glandular 
trichomes are found in Q. salicifolia. 


1104 BRIT.ORG/SIDA 22(2) 


Trichome morphology and species identification 

There are several instances where trichome morphology can be a valuable tool in the 
identification of red oak species, especially of those with similar overall features. For ex- 
ample, Q. crassipes and Q. mexicana (Figs. 7A, B) are frequently confused because they 
have similar leaf morphology. However, examination of trichomes on the abaxial leaf 
surfaces ie on ae fferences: while Q. crassipes has fasciculate stipitate trichomes, Q. 


mexic ile contorted trichomes. Reproductive characters (e.g. fruit 


morphology) agree with their distinctiveness based on trichome morphology. Another 
example of the utility of trichome morphology for species identification is found in the 
Crassifoliae complex (sensu Vazquez 2001, which consists of fourteen species charac- 
terized by the presence of a pale-yellow indumentum on the abaxial leaf surface, and by 
a morphological resemblance to leaves of Q. crassifolia. Two species often confused with 
Q. crassifolia are Q. mcvaughiiand Q. fulva. While the abaxial leaf surface of Q. crassifolia 
has distinctive lfasciculate stipitate trichomes, Q. mcvaughii displays fasciculate sessile 
trichomes and Q. fulva shows multiradiate trichomes on their abaxial leaf surfaces. In 
fact, trichome differences between Q. crassifolia and Q. mevaughii, together with other 
morphological and reproductive characters were used in the segregation of these mor- 
phologically similar species (Spellenberg 1992). 

A potential problem in trichome classification is that there are cases when there is 
not a clear-cut distinction among types. One case involves Hardin’s (1976) glandular 
rosulate trichome type, characterized by the rosette pattern of the rays and, most impor- 
tantly, the rays having a thin cell wall. In practice, it is difficult to determine the nature 
of the cell wall unless ultrastructural studies are carried out. In the absence of such stud- 
ies and taking into account ray arrangement, these trichomes should be classified as 
multiradiate. In this study, all trichomes with rays originating from a single point and 
diverging in different directions were considered within the multiradiate type 

A second problem in the identification of trichome types is the transition of one 
type into another. In the specimens examined, morphological transitions were observed 
between fasciculate sessile and fasciculate stipitate trichomes, and from fasciculate sessile 
to stellate. These examples agree with Jones’ (1986) statement that sometimes there are 
no discrete boundaries between trichome types and that “.. most forms intergrade with at 
least one other type...” 

Despite these potential limitations, SEM has been a key factor in the discovery of 
three new trichome types, as well as in the finding of three trichome types not previ- 
ously reported for red oaks. Furthermore, study of trichomes under SEM has revealed key 
morphological differences in apparently similar trichome types. Detailed examination 
of trichomes from Mexican oak taxa revealed that four different trichome types had been 
included under the “starred” category in the taxonomic literature. The trichome comple- 
ments of mature leaves are useful in the identification of morphologically similar oak 
species when taken together with other vegetative and reproductive features, as indicated 
in the dichotomous key below. This work PEpyinee a standard terminology that could be 


used in future taxonomic treatments to aid in sf : iption of Mexican oak taxa. 


= 


DICHOTOMOUS KEY FOR IDENTIFICATION OF MEXICAN RED OAK SPECIES 
. Abaxial leaf surface either completely devoid of trichomes or appearing glabrous except for 
scattered glandular or nonglandular trichomes. 
o suborbicular or pandurate; abaxial leaf surface with abundant simple 


2. Leaf shape ovate 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 


eink: 2 
Wier 


year 
; KE hs ea 

BLEUE 
. o: + 


(a 


ict the abaxial leaf surface, X 74.C.—D. Q. mexicana 
L ae oe | _ m4 «74, 


Fic.7. A.—B. Quercus crassipes, left i i i 
F y TIGH, I 
+ | H + €. H lo¢ H | = 


uniseriate glandular trichomes and a few fasciculate stipitate trichomes restricted to the 
Q. coccolobifolia 


secondary vein axils 
2. Leaf shape elliptic, lanceolate, ovate-lanceolate, or oblanceolate; abaxial leaf surface devoid 


of trichomes or with scattered stellate or multiradiate trichomes. 
3. Leaf margin entire and undulate 
4. Leaves 4-6 times long as wide; abaxial leaf surface with sparse multiple stellate tri- 
chomes or multiradiate trichomes distributed throughout the lamina and vein axils; 
fruits with biennial maturation. 
5. Secondary veins 10-15; two-thirds of the acorn included in the cup; epidermis on 


Q. gentryi 


the abaxial leaf surface bullate 
5. Secondary veins 15-25; one-third of the acorn included in the cup; epidermis on 
Q. salicifolia 


the abaxial leaf surface smooth 
Leaves 2—2.5 times long as wide, abaxial leaf surface lacking trichomes on the lamina 


but with fasciculate stipitate trichomes on the vein axils; fruits with annual maturation 
Q.acherdophylla 


aN 


1106 BRIT.ORG/SIDA 22(2) 
3. Leaf margin entire and straight or with short teeth. 


6. Leaf base attenuate, cuneate or decurrent; mature leaves 3.6-4.2 times long as wide 


Q. affinis 
6. Leaf base round or obtuse; mature leaves 2.4—2.6 long as wid Q. depressa 


“Abn leaf surface with abundant and persistent nonglandular trichomes, sometimes trichomes 


7. Abaxial leaf surfac 


e sparingly pubescent or glabrescent; if sparingly pubescent, at low mag- 
nifications the epidermis could be seen through the trichomes 
8. Trichomes on axial leaf surface have a punctat ey trichomes fasciculate con- 


9. Leaves elliptic-oblong, always entire, veins 6-16 
9. Leaves elliptic-lanceolate, elliptic-oblanceolate, 
oothed, veins 5-8 


Q. mexicana 


ometimes obovate or ovate, entire or 


Q. eduardii 


Trichomes on abaxial leaf surface have a floccose or arachnoid appearance; trichomes 
ion stipitate, fasciculate sessile simple, or multiradiat 
es orbicular, orbicular-pandurate, or broadly es twigs 4-8 mm thick. 


(ae leaves with 4-8 teeth; infructescence up to 6 mm long bearing 1-2 fruits 


Q. tarahumara 
11. Mature ik with 13-19 teeth; infructescence from 4.7 to 15 cm long bearing 
3-9 fru Q. radiata 
10. Leaves sii ,lanceolate, ovate or obovate; twigs 0.75-4(-5) mm thick. 
2. Fruit cupule with the margin involute; cupule diameter 20-28 mm; secondary 
veins 10-19 


Q. planipocula 
2. Fruit cupule with the margin straight; cupule diameter 7-15 mm;secondary veins 
= 12, 


13. Fruits with annual maturation; bullate epidermis on the abaxial leaf surface; 
secondary veins 5-12 


14. Leaves obovate to elliptic; trichomes on the abaxial leaf surface fascicu- 
late sessile or fasciculate shortly stipitate; leaf margin toothed; distributed 
y SUIp C 
in the Sierra Madre Oriental 


. hirtifolia 
4. Leaves elliptic-lanceolate, elliptic-oblanceolate, sometimes obovate or 
ovate; trichomes on the abaxial leaf surface fasciculate distinctly stipitate; 
leaf margin entire or entire-aristate; distributed in the Sierra Madre 
Occidental Q. aristata 
13. Fruits with biennial mat th epidermi the abaxial leaf surface; 
secondary veins 4-5 
15. Aba 


xial leaf siintack appearing glabrous but with two types of glandular 
nontrichomes visible at 10:small multiradiate of short rays distributed 


on the abaxial lamina and fasciculate stipitate trichomes confined 
to the secondary vein axils 


Q. hintoniorum 
15. Abaxial leaf surface pubescent but the trichomes detach with age; tri- 


chomes fasciculate sessile simple with long intertwined rays distributed 
enn e the lamina 


Q. hypoxantha 
7. Abaxial leaf surface densely aie the epidermis completely covered with trichomes. 
16. Leaves elliptic, lanceolate, obla | e. 


17. Leaf margin always en a arista 


18. Abaxial leaf surface Suess eee cent, the indumentum whitish, trichomes 
multiradiate or | 


t appearing fasciculate sessile at 10x: ;apex acute 


9. Leaf margin flat; mature leaves 2.6-3 times long as wide; petiole 2-6 mm 
g 


lon Q. durifolia 
19. Leaf margin revolute; mature leaves 3.75—5 times long as wide; petiole 5-15 
mm long hypoleucoides 
18. 


Q. 
Abaxial leaf surface loosely pubescent, the indumentum pale yellow, trichomes 
clearly fasciculate stipitate at 10x; apex rounded oro 


obtuse 
20. Mature leaves about 2.5 times longer than wide; cupbie hemispheric or tur- 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 


17. Leaf margin mainly toothed, rarely entire with aris 


16. Leaves obovate, suborbicular, nes or very broadly elliptic. 
25. The indumentu 


25. Thei 


1107 


binate, the cupule margin not involute; 
sessile and fasciculate stipitate trichomes 
20. Mature leaves 4—-4.5 times longer than wide; cupule hemispheric with invo- 


lute margin; current year twigs with fasciculate sessile and multiradiate tri- 
chomes 


year twigs with fasciculate 
dysophylla 


Q. crassipes 


21. Twigs 3-5 mm thick; abaxial leaf surface with ake stipitate trichomes;cup 
very shallow up to 5 mm tall, enclosing 1/4 of the nut 
21. Twigs 1-3 mm thick; avant leaf surface with Tescicilane ee simple, 
multiradiate, or fasciculat rested trict 
- enclosing 1/3 to 1/2 of the n 
2. Abaxial leaf surface with ee sessile trichomes of 6 to 10 long inter- 
twined rays. 
23. Petioles 3-5(—7) mm long; fasciculate sessile trichomes twisted at the 
e and detaching with age; leaf margin slightly revolute Q. coahuilensis 
23. Petioles 5-25 mm long; fasciculate sessile trichomes not twisted at the 
base and pocisting with age; leaf margin flat 
22. Abaxial leaf surface with multiradiate ee lat il 
of 20-22 short rays. 


Q. hintonii 


’ 


a 


Q. castanea 


omes 


24. Leaf base decurrent or cuneate; leaf tapering toward the apex; leaf mar- 
gin not revolute Q. scytophylla. 
24. Leaf base rounded, subcordate or cordate; leaf apex acute or obtuse; 
leaf margin revolut 


Q. sideroxyla 


n the abaxial leaf surfaces detaches with age; twigs 0.75- 
thick; petioles eae fegaleh 

indumentum on the f surf 

petioles densely pubescent pubescent or - glabrescent, 
26. Petioles 0.4-25 mm long; cup 7-9 mm wide, 4-5 mm tall; acorn 7-11 mm long 


o hypoxantha 
hed;twigs 2-13 mm thick; 


m wide Q. conzattii 
26. Petioles 25-37 mm long; cup 8-24 mm wide, 5-15 mm tall; acorn 10-24 mm 
long, 7-21 mm wide. 
27. Twigs fines season 8-13 mm thick; infructescences 3.5 to 10 cm long bear- 
ing 6-10 Q. urbanii 


27. Twigs ee season 2.5-6.0 mm thick; infructescences less than 2 cm long 

bearing 1-2 fruits 
28. indumentum on the apaalal leaf surface white or pale yer, leaves 
with 1 teeth; sp glandular trichomes on the abaxial leaf surface 


Q.candicans 
28. Indumentum on the abaxial leaf surface yellow or yellow-brownish; 
leaves with 1-11 teeth; abundant glandular trichomes on the abaxial 
leaf surface. 
29. Abaxial leaf surface with a felted indumentum composed of 
multiradiate trichomes with straight rays; rim of cupule involute 


Q. fulva 


29. Abaxial leaf surface with a woolly indumentum composed of fas- 
ciculate trichomes with long tortuous rays; rim of cupule straight. 
30. Mature leaves deciduous; trichomes on the abaxial leaf surface 
ee stipitat Q. crassifolia 
30. Mature leaves Se Serre on the abaxial leaf surface 
fasciculate sessile simple 


Q.mcvaughii 


1108 BRIT.ORG/SIDA 22 


2) 
ACKNOWLEDGMENTS 


| wish to thank Kevin Nixon for sharing his knowledge on oak systematics, the Bailey 
Hortorium, Department of Plant Biology, at Cornell University, for financial assistance 
for the completion of this research, Richard Spellenberg for his great suggestions for the 
improvement of this manuscript, Brian Phillips and Amy McEuen for their comments 
on the manuscript and testing the dichotomous key, and Jeff Bacon, Hector Vazquez 
Villagran, Liliana Vazquez Villagran,and Rebeca Ocampo for their assistance in the field. 
Lalso want to thank Richard J. Jensen (NDG) and Paul S. Manos (DUKE) for their helpful 
suggestions as reviewers for Sida. 


REFERENCES 

Bacon, J.R.and R. Speccensers. 1996. Hybridization in two distantly related Mexican black oaks Quercus 
conzattii and Quercus eduardii (Fagaceae: Quercus: Section Lobatae). Sida 17:17-41 

Brito, M.A. and J.N. Labat. 1987. Los encinos (Quercus) del Estado de Michoacan, México. Centre 
d'Etudes Mexicaines et Centroamericaines. SARH-INIF. Mexico. 

Bussorti, FP. and P. Grossoni. 1997. European and mediterranean oaks (Quercus L.; Fagaceae): SEM 
characterization of the micromorphology of the abaxial leaf surface. Bot J Linn Soc 124:183-199. 

Camus, A. 1934-1938.Monographie du genre Quercus. Vol 1.Sous-genre Cyclobalanopsis. Sous-genre 
nee sections Cerris et Mesobalanus. Academie des Sciences. Lechevalier, Paris. 

Camus, A. 1936. Monographie du genre Quercus. Vol 2 Les chenes Section Lepidobalanus. Academie 
des en Lechevalier, Paris. 

Camus, A. 1952-1953. Monographie du genre Quercus. Vol. 3. Sous-genre Quercus. Sections 
Protobalanus et Erythrobalanus. Academie des Sciences Lechevalier, Paris. 

Dyat, S.A. 1936.A key to species of oaks of eastern North America based on foliage and twig charac- 
ters. Rhodora 38:53-63. 

GeLuNi, R.F., BD. Bussorm, P. Bertini, P. Grossoni, and A. Bor tacci. 1992. Species of the genus Quercus in Italy: 
characterization by means of leaf surface observation. Giorn. Bot. Ital. 126:481-504 

Gonzalez, L.M. 1986. Contribuci6n al conocimiento del género Quercus (Fagaceae) en el estado de 
Jalisco. Instituto de Botanica. Universidad Autonoma de Guadalajara. Mexico 

Haroin, JW. 1976. Terminology and classification of Quercus trichomes. J Elisha Mitchell Sci Soc 92: 
151-161. 

Harbin, J.W. 1979a. Patterns of variation in foliar trichomes of eastern North American Quercus.Amer. 
J.Bot.66:576-585. 

Harbin, J.W. 1979b. Atlas of foliar surface features in woody plants, |. Vestiture and trichome types of 
eastern North American Quercus. Bull Torrey Bot Club 106:313-325 

INAMDAR, J.A. AND N.V. Rao. 1983. Light and scanning electron microscopic studies on trichomes of 
some Brassicaceae. Feddes Repert. 94:183-190. 

JENSEN, R. 1997. Quercus section Lobatae.|n: Flora of North America Editorial Committee, eds, Fl.North 
Amer. 3:445-506 Oxford Univ. Press, New York and Oxford. 

Jones, H. 1986. Evolution of the Fagaceae: The implications of foliar features. Ann. Missouri Bot. Gard. 
73:228-275. 

Kissuine, P.1977.Les poils des quatre especes de chenes du Jura (Quercus pubescens, Q. petraea, Q. 
robur et Q. cerris). Ber Schweiz Bot Ges 87:1-18. 

Kissuin, P. 1993. Heteroindumentum: Des poils de taxons differents sur la meme plante. Bot Helv 
103:83-99, 

Liamas. F., C. Perez-Morates, A. Acebo, and A. Penas. 1995. Foliar trichomes of the evergreen and 


VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1109 


semideciduous species of the genus Quercus (Fagaceae) in the Iberian Peninsula. Bot.J.Linn.Soc. 
117:47-57, 

Manos, PS. 1993a.Cladistic analyses of molecular variation of“higher” Hamamelididae and Fagaceae, 
and systematics of Quercus section Protobalanus. Ph.D. diss. Cornell University, Ithaca, NY. 

Manos, PS. 1993b. Foliar trichome variation in Quercus section Protobalanus (Fagaceae). Sida 15: 
391-403 

Manos, P.S., Nixon. K.C., Doyte, J.J. 1999. Phylogeny, biogeography, and processes of molecular differ- 
entiation in Quercus subgenus Quercus (Fagaceae). Molec. Phyl. Evol. 12:333-349. 

McVaucH, R. 1974. Flora Novo-Galiciana, Contr. Univ. Mich. Herb. 12(1):1-43. 

Metcatre, C.R. AND L. CHALK. 1950. Anatomy of dicotyledons. Vol 2. Oxford. 

Nixon, K.C. 1984. A biosystematic study of Quercus series Virentes (the live oaks) with phylogenetic 
analyses of Fagales, Fagaceae and Quercus. Ph.D. Dissertation.The University of Texas at Austin, Texas. 

Nixon, K.C. 1993. The genus Quercus in Mexico. In: Ramamoorty, TP et al., eds. Biological diversity of 
Mexico: origins and distribution. Oxford Univ. Press. Pp. 447-458. 

Nixon, K.C. and K.P. Steete. 1981. A new species of Quercus from Southern California. Madrono 28: 
210-219. 

Nixon, K.C.and C.H.MUuLver. 1993. The Quer: | (6 tl on \plex (Fagaceae) in northeastern Mexico. 
Brittonia 45:146-153. 

Otsson, U.1976.The structure of stellate trichomes and their taxonomic implication in some Quercus 
species (Fagaceae). Bot. Notiser. 128:412-424 

Oostuuizen, L. 1983. The taxonomic value of trichomes in Pelargonium (Geraniaceae). J. S. Afr. Bot. 
49:221-242. 

Romero, S. 1993. El género Quercus (Fagaceae) en el estado de México. Master's thesis. Universidad 
Nacional Auténoma de México. Mexico 

Rzepowski, J. 1965. Relaciones geograficas y posibles origenes de la flora de México. Bol. Soc. Bot. Méx. 
29:121-177. 

Rzepowski, J. and G.C. Rzevowski. 1979. Flora fanerogamica del Valle de México. CECSA, México. 

Sarou, O. AND M. Saint-Maatin. 1989. Foliar trichomes of some perimediterranean oaks. Bull. Soc. Bot. 
France Lett. Bot. 136:291-304. 

SHANMUKHA, Rao S.R. 1987. Structure, distribution and classification of plant trichomes in relation to 
taxonomy: Sterculiaceae. Feddes Repert. 98 (1-2):127-135. 

SPELLENBERG, R. 1992. A new species of black oak (Quercus, subg. Erythrobalanus, Fagaceae) from the 
Sierra Madre Occidental, Mexico. Amer. J. Bot. 79:1200-1206. 

SPELLENBERG, R. 1998. Quercus lesueuri, un miembro meridional del complejo Q._ undulata (Fagaceae, 
subgenero Quercus). Acta Bot. Mexicana 42:25-33. 

SpeLLENBERG, R. 2001. Taxonomic overview of the oaks of La Frontera. In:G.L.Webster and C.J. Bahre, 
Changing Plant Life of La Frontera, Univ. New Mexico Press, Albuquerque. Pp. 195-211. 

SPELLENBERG, R., J.R. BACON, AND M.S. GonzALez. 1998. Los encinos (Quercus, Fagaceae) en un transecto 
sobre la Sierra Madre Occidental. Boletin Instituto Botanica, Universidad de Guadalajara, Mexico 
9:357-38/7. 

THomson, P.M. and R.H. Monvensrock. 1979. Foliar trichomes of Quercus subgenus Quercus in the East- 
ern United States. J. Arnold Arbor. 60:350-367. 

Tucker, JM. 1952. Evolution of the California oak Quercus alvordiana. Evolution 6:162-180. 

Tucker, JM.and C.H.Mutter. 1957.A reevaluation of the derivation of Quercus margaretta from Quercus 
gambelii. Evolution 12 -17. 

Upnor, J.C. 1962. Plant hairs. Gebrtider Borntraeger. Berlin-Nikolassee. 


a 


1110 BRIT.ORG/SIDA 22(2) 


tadn da CGiier 
DtdGO Qe WUE! 


VALENCIA, S. 1989.Contribucidn al conocimiento del género Quercus (Fagaceae) | 
rero, México. Undergraduate thesis. Universidad Nacional Aut6noma de México. 

VAzquez, M.L. 1992. El género Quercus en el estado de Puebla, Mexico. Undergraduate thesis. Univer- 
sidad Nacional Autonoma de Mexico. 

VAzauez, M.L. 2000. Flora del Valle de Tehuacan-Cuicatlan. Fagaceae. Instituto de Biologia, Univer- 
sidad Nacional Autonoma de México, Fasciculo 28. Mexico. 

VaAzauez,M.L. 2001 Molecular and morphological studies on Mexican red oaks (Quercus sect.Lobatae). 
Ph D. Dissertation. Cornell University, New York. 


THOMAS WALTER TYPIFICATION PROJECT, L. 
OBSERVATIONS ON THE JOHN FRASER FOLIO 


Daniel B.Ward 


Department or Borany 
I fap #C]/ 
UTIIVET OF ity GIT lorida 


Gainesville, Florida 32611, U.S.A. 


ABSTRACT 


The Natural History Museum, London, is the repository of a large volume containing plants collected by John 
Fraser in the southeastern United States in the 1780s and commonly known as the : ANalter Herbarium.” The 


importance of this collection is that mony a its ceed were seen and petheps ued Py Pom as Walter, au- 
thor of Flora Caroliniana, the first A | pt Linnae an | t Ade escription 


is provided here of this folio and the collection it contains. 


RESUMEN 


El Natural History Museum de Londre | | | 
John Fraser en el sudeste de los Estados ee en los 1780s y cenesido seeaalnente como el‘ “Walter He ee 


La importancia de esta coleccion radical en que muchos de sus especimenes fueros vistos y quizas usados por 


Thomas Walter, autor de la Flora micas aa: la ae flora americana en adoptar la oe Linneana y 
la nomenclatura binomial. Se hace un le este volumen y la coleccion que contiene 


The Thomas Walter Typification Project is the term now given of an effort, extended in- 
termittently over forty years, to bring understanding and nomenclatural precision to the 
many plant names published by Thomas Walter in his pioneer Flora Caroliniana (1788). 
Walter, an English rice-plantation owner and amateur but skilled botanist, resident of 
South Carolina, was the first to prepare a flora of American plants utilizing Linnaeus’s 
binomial nomenclature and sexual system of classification. A brief but close relation- 

developed between Walter and John Fraser, a Scottish plant explorer who in the late 
1780s gathered numerous specimens from the southeastern United States into a large 
folio herbarium now held by the Natural History Museum, London. The specimens of 
this herbarium were seen and may in part have been used by Walter in preparation of the 
diagnoses of his new species. The present report is an account of the physical character- 
istics of this historic collection. 

The information of this report is derived from notes taken during a two-week ex- 
amination of the folio by the present author in July 1984, and from 8x10 glossy photos 
enlarged from negatives made of the folio by Dr. Bernice G. Schubert in November and 
December 1946 (Fernald & Schubert 1948). (Commercial microfiches distributed in 1985 
were of lesser clarity. Digital images prepared in 2005 were not yet available.) Though 
much remains unknown, many aspects of the interrelated lives and works of Thomas 
Walter and John Fraser have been documented by Britten (1921), Coker (1910), Hogg (1852), 
Maxon (1936), Rembert (1980), Simpson et al. (1997), Small (1935), and Ward (1962 1977). 

The specimens are mounted on the pages of a large book, a folio, 54.5 cm tall, 38 cm 
wide, about 8 cm thick, bound in green cloth and half-leather with 5 transverse raised 
bands and several thin gold lines. The pages are 52 cm by 35.5 cm, of heavy linen-rag 
watermarked paper, bound permanently along a lateral margin. The book is of commer- 


SIDA 22(2): 111- 118.2006 2006 


1112 BRIT.ORG/SIDA 22(2) 


cial origin, manufactured in advance of its present use; it was not made by binding to- 
gether separate sheets to which the plants had been attached. Two initial pages are un- 
numbered; each subsequent page has been numbered by hand near its upper margin. 
The folio consists of 119 numbered pages (not 117, as reported); two numbers (61 and 62) 
were used twice. 

The frontispiece, the unnumbered first page of the folio, bears an attached sheet (right- 
diagonals indicate new lines; left diagonals reflect superscripted letters): “The Herbarium 
of Tho\s Walter Esq. of South Carolina. / Author of the Flora Caroliniana, pub. 1788 / 
Presented May 23, 1849, to the / Linnean Society of London, by John Fraser, / son of John 
Fraser, the indefatigable / North American Botanical Collector, / from the years 1786 to 
I811. He died / in London, the latter year.” This slip (as believed by James Britten 1921) is 
in the hand of Fraser’s son, John Fraser fil. A second, unknown writer has restated these 
words by inscription onto the page itself: “The Herbarium / of Thomas Walter / author 
of the / “Flora Caroliniana.” / Presented to the Linnean Society, 23 May 1849, / and pur- 
chased at the sale of the Society’s / Surplus Collections in 1863 (for 15/.).’ The frontispiece 
also bears a detailed listing (probably by Britten) of the persons, into the 20th century, 
whose annotations are to be found within the folio. 

The title page of the folio, also unnumbered, bears the boldly written words, “Walter's 
Herbarium,” the dates “1786-1788,” and the initials, “JE” The hand is that of John Fraser. 
The dates encompass the frenetic eighteen months—from September 1786 until March 
1788—during which Fraser traveled and collected in the American Southeast. 

The plants of the collection are mounted (counterintuitively) on the left-hand pages 
of the opened folio. The herbarium consists of 690 specimens (of which three are repre- 
sented only by their labels and by stains on the page to which they had been attached), 
The specimens are mounted with as few as 1 and as many as 12 ona single page (an aver- 
age of 5.8 specimens per page). 

For clarity of reference in present and future study, images (xeroxes from photos) of 
the specimens have been enumerated by two-part designators: each individual specimen 
is given the number of the page on which it is mounted, followed by a capital letter. (The 
two pages with duplicated numbering, p. 61 and p. ©2, are distinguished by suffixes “a” 
and “b.”) The specimens, insofar as their arrangement on the page permits, have been 
assigned a designator in alphabetical order, from upper left to upper right, second-row 
left to second-row right, down the page. A copy of the images showing the assigned des- 
ignators has been provided to the Natural History Museum, for placement with the folio. 
These designators are suggested for use in specimen citation, to replace the use of page 
numbers or other accompanying numbers seen on many labels, as resorted to by other 
authors. The designators are employed in the present discussion where appropriate. 

Nearly all specimens are accompanied by a small label. (A very few have no label; a 
few have either two labels, or a label clearly belonging with another specimen.) The la- 
bels vary in proportion and dimension, but are mostly of horizontal rectangles, 5 to 8 cm 
long, 2 to5 cm high. All have clearly been individually and somewhat roughly cut from 
other sheets. A few (e.g., 41-E, 51-A, 65-A, 71-A, 80-A, 83-A, 83-E) have been folded trans- 
versely, the lower half tucked behind the upper half which bears the writing. Two dis- 
tinct paper stocks were used for the labels, both of “laid paper,” formed froma linen slurry: 
the one thin, tan, with laid-lines spaced +9 per cm; the other heavier, whiter, with laid- 
lines +7 per cm. 

Most of the specimens are attached to the label by one slit or more often two parallel 


— 


— 


R IT COLLECTION 1113 


slits cut in the label, with the plant stem passed under the resulting strap. At times the 
slits are in the form of shallow V’s (e.g., spms. 24-B, 31-C, 32-C, 34-B, 6la-F), indicating 
they were cut by folding the label, then making the two cuts with two snips of a scissors. 
Frequently part of the writing on the slip is beneath part of the specimen, obviously 
having been inscribed prior to attachment of the plant. A few of the specimens have been 
attached to the label by slender pins (e.g., 46-B, 60-E 61la-A, 92-J, 105-C). Others have been 
given firmer mounting, either to the label or to the underlying sheet, by conventional 
narrow tape strips. 

A significant achievement of the Project has been the determination of nearly all of 
the handwritings on the labels. Yet, for reasons explained elsewhere (Ward, in press), this 
information is of less value than might be assumed for determination of the collector of 
the accompanying specimen or for prediction of its use by Thomas Walter in forming the 
diagnoses of his Flora. 

Two handwritings predominate on the labels: Thomas Walter (ca. 1740-1789), and 
John Fraser (1750-1811). The hand of Fraser's son, John Fraser (1780-1861) [not “1799- 
1860?”—Fernald & Schubert 1948] also appears to be present, though rarely. External evi- 
dence of these hands is provided by a 158-word letter written by Walter in 1783 and re- 
produced by Ewan (1979), and by a 48-word letter written by Fraser’s son in 1818 in the 
Autograph Collection of the Natural History Museum. Fraser's hand is established by the 
few words and dates of the title page and by occasional brief, initialed notes accompany- 
ing the specimens (e.g., 67-A, 78-E). The third hand probably of Fraser fil. differs mark- 
edly from that of Fraser and also somewhat from that of Walter. tis found on a few labels 
(ge GF 26) 676 lbp 105-G, 115-G). Numbering of the pages of the folio 
may also have been by Fraser fil; these numbers are not in the hand of Fraser sr. 

Once determination was made that two writers were responsible for the great major- 
ity of the labels, most of the handwriting could be assigned. Past authors have referred in 
general terms to similarities with that of Walter, Fraser, or Fraser's son, but few authors 
have claimed which hands were found in association with specific specimens, and none 
are known to have based nomenclatural arguments on such identifications. [Blake 
(1915:130) correctly identified Walter's hand on four labels, but wrongly concluded “most 
of the remaining labels are in another hand, which from its agreement with some writ- 
ing on the title page of the volume may be affirmed with considerable certainty to be 
that of John Fraser himself | Yet close—and tediously prolonged—examination of the hand- 
writing of the labels and of the available handwriting samples does usually permit cer- 
tain identification 

John Fraser’s hand in most cases is unambiguous. It is a smoothly flowing script, 
with the letters consistently formed and widely spaced. His capital “A” is of the classic 
“inverted V” and uniformly shows the cross-bar sweeping far left across the ascent. His 
capital “C” is always shaped with both its beginning and terminus well to the left of the 
figure. His capital “P” is initiated by a strong upstroke, sharply reversed to a downstroke, 
again reversed to form a clockwise loop. Many of his other capital letters also show dis- 
tinctive features, though his lower case letters are largely of standard form. He is unfail- 
ing in his practice of using capitals for the initial letters of both genera and epithets. His 
hand is of an extrovert—if this term can be applied toa writing style—fitting the man of 
confidence and courage his travels and ventures show him to have been. 

Thomas Walter’s hand is far more cautious, cramped, and poorly, unevenly formed. 
His capital “A” is at times an “inverted V,” but his cross-bar is formed separately and 


1114 BRIT.ORG/SIDA 22(2) 


scarcely touches the ascent; for many of his plant names his capital “A” is merely lower 
case writ large. His capital “C” is of the same form as Fraser's, but only the beginning is 
from the left; the lower terminus invariably links with the next letter. His capital “P” is of 
a smooth clockwise swirl, with the initial stroke directed downward. With few excep- 
tions his epithets are not capitalized. His hand in general is that of a busy, quick thinking 
person, not overly concerned with the niceties of precise letter formation; in modern days 
his script might have been that of a doctor. 

A readily available sample of the two handwritings is given in photographs shown 
by Faden (1989:46-47). There, the single word “Commelina” is written on the label of a 
specimen of C. erecta by Walter, and on the label of a specimen of C. caroliniana by Fraser. 
The Walter label also bears “305,” a number written by Fraser. 

John Fraser fil’s hand is poorly understood because of its infrequent appearance. 
Though it is unlikely to be mistaken for that of John Fraser sr, brief sequences of letters 
(as in plant names or brief notes) may not show marked contrast with that of Walter. A 
character that may be useful is the “square r” here attributed to Fraser fil., while the lower- 

case “r” of both Fraser sr. and Walter is usually an abrupt simple upward pip 

More than half of the labels (371, or 54%) bear three-digit (rarely two- digit) numbers 
ina darker ink anda coarser pen, and seem to have been written prior to other markings. 
The hand forming these numbers is that of John Fraser; his distinctively malformed “8” 
appears also as part of the date given on the title page of the folio. (Representative speci- 
mens so marked: 1-H, 27-C, 34-D, 51-B,54-D, 59-E, 66-C, 101-C, 103-F 107-A.) Britten (1921) 
was unable to discover the use to which Fraser put these numbers, nor have others sug- 
gested any logical purpose. The numbers correspond neither to the present arrangement 
of the specimens nor to the sequence of Walter's Flora (nor presumably to its parent manu- 
script). Many of these numbers have been struck by a single diagonal slash; most of the 
rest are accompanied by a small checkmark. No numbers are repeated. The numbers may 
differ in size between two adjacent specimens (e.g.,62a-B and 62a-C, 94-D and 94-E), dem- 
onstrating that specimens now adjacent in the herbarium were numbered at different 
times. Frequently the numbers are partially cut away at edge of the label (e.g, 38-1, 50-E, 
80-D, 84-A, 86-C, 92-A, 105-B, 108-F). These characteristics support the speculation that 
Fraser's bers served as field references, perhaps coupled toa now-lost separate record 
of source and date, the pages later cut to their present dimensions and used as permanent 

labels. The few two-digit numbers of this series (six exceptions, but not clearly in the 
same hand), together with knowledge that Fraser spent earlier years (1780-1784) in New- 
foundland where he also studied plants (Fraser 1789; Hogg 1852), suggests that these num- 
bers may have been a continuing enumeration of his collections begun before he first 
came to the Carolinas. Certainly, aside from their initial purpose, their presence on the 
label indicates that the specimen is of Fraser's collection, whether or not it was later seen 
by Thomas Walter 

Another series of numbers seems not to have been noted. Each number of this small 
class (20 numbers) is preceded by a capital “F” or “FC.” All are clear references to the num- 
bered genera in Flora Caroliniana. Examples: L00-C is named “Staphylea Trifolia” and 
numbered “F 132,” while genus 132 is Staphylea; 106-A is named “Verbena Urticifolia” 
and numbered “F 236,” while genus 236 is Verbena with the single species V. urticifolia; 
116-Cand 116-D both bear the name “Syntherisma” and are numbered “F 35,” while genus 
35is Syntherisma (= Digitaria). Further examples: 32-A, 32-B, 32-C, 32-D, 38-], 48-C, 115- 
G, 117-A, 117-D, 117-E.) Most of the numbers occur in clusters (e.g., 32-A thru 32-D, 101-A 


1115 


thru 101-G, 115-G thru 117-E), indicating they were assigned after the specimens were 
arranged in the herbarium. The hand inscribing this series of numbers is probably that 
of Fraser (e.g., 38-J, 48-C, 100-C), though at times the “F” seems unlike his. Possibly it is 
that of his son; it is not that of Walter. These numbers were certainly assigned after pub- 
lication of Flora Caroliniana, or at least after access to the completed manuscript. This 
second series of numbers appears to have been a partial, never completed attempt to match 
the specimens collected by Fraser to the appropriate souls as Su blal aice by Walter. 

Within the folio the specimens are arranged in l sequence, as determined 
by the names borne by the original labels. These names, in turn, are largely those used in 
Walter's Flora Caroliniana(1788).[The few exceptions are usually in the hand of John Fraser 
and beara name from Linnaeus that was not used by Walter (e.g., 37-G, “Cornus canadensis’; 
52-E, “Globularia nudicaulis”, 58-C, “Helonias bullata”), or a comment by Fraser or Walter 
in place of a name le.g., 38-D, “Corypha arborea” (= Sabal palmetto), 43-B, “Erythronium 
Dens Canis” (= E.americanum),55-C, “Hedysarum Flore magnus’ (= Desmodium cuspidatum), 
67-D, “Lupinus affinis” (= Crotalaria rotundifolia), 72-A, “Mespilus Large Cock Spur’ (= 
Crataegus crus-galli); 98-D, “Smilax Baccis albidis” (= Smilax laurifolia)], 

Of the specimens bearing an identification on the label (either complete, or only to 
genus), 345 (50.1%) bear Walter's hand, 230 (33.4%) bear Fraser's. Often the hands of Walter 
and of Fraser are found on the same label. On 49 labels Walter identified the specimen to 
genus and Fraser completed the naming by adding the species. On none of the labels did 
Fraser identify the specimen to genus, with Walter adding the species. 

Handwritings by other persons have been added over the years, usually in the form 
of identifications. They may be on separate slips, on the adjacent sheet, or on the original 
label itself. Those writers identified on the frontispiece of the folio (by James Britten) are: 
Oakes Ames, Sidney F Blake, Nathaniel Lord Britton, Alva A. Eaton, Asa Gray, Albert S. 
Hitchcock, Charles S. Sargent, James E. Smith, and Anna M. Vail. Others include James 
Britten, James E. Dandy, Merritt Lyndon Fernald, William T. Gillis, A.J. Kosterman, John 
Lewis, Mildred EF. Mathias, and Charles A. Weatherby. More recent annotations (since 1983) 
are by Joseph Ewan, David K. Northington, David H. Rembert, Daniel B. Ward, and others. 

Identification of the specimens is no easy task. As noted in 1839 by Asa Gray (J.L. 
Gray 1893), many of the specimens are “mere bits,” small vegetative fragments, single 
leaves or single flowers, often wholly unidentifiable from photographs and scarcely less 
so by direct examination without painstaking comparison to known materials. Signifi- 
cant efforts to identify more than single genera by direct examination are few. Hitchcock 
(1905) reviewed Walter’s grasses and matched many of their names with Fraser's speci- 
mens. Blake (1915) discussed 22 Walter names and their corresponding specimens and 
made new combinations where indicated. Fernald & Schubert (1948), working from pho- 
tographs, identified—not always correctly—19 specimens in the herbarium. Others have 
examined the species of individual genera and published their conclusions. Still others, 
notably Oakes Ames with the orchids, and Asa Gray and David H. Rembert with large 
parts of the entire herbarium, have given significant time to identifying the specimens, 
but have left no record other than their annotations and notes. 

At the present writing perhaps one third of the specimens has been given a credit- 
able identification, one third a tentative judgment, and one third no more thana cautious 
assignment of genus. Complete identifications must await opportunity for study by per- 
sons who have simultaneous access to the Fraser specimens and to a convenient refer- 
ence collection. 


1116 BRIT.ORG/SIDA 22(2) 


Only two of the 690 specimens bear a date on the original label. Spm. 78-E, probably 
Platanthera flava, is atypically instructive: “Orchis / found near the / Table Mountains / 
llth June / 1787 /JE” The hand, confirmed by the initials, is that of John Fraser. Spm. 100- 
F, Gillenia trifoliata, is briefly informative though lacking the year of collection: “16 June 
/ Indian Emetic,” written by Fraser, followed by “Spiraea trifoliata,” written by Walter. 

Equally few specimens bear a geographic source on the original label. Only two la- 
bels give locations; both are in Fraser's hand. As noted, spm. 78-E, probably Platanthera 
flava, was collected near the “Table Mountains,” perhaps Table Rock, Pickens Co., South 
Carolina (less likely: Table Rock, Burke Co., North Carolina). Spm. 67-A, Lupinus villosus, 
was labeled “Blue Flowering / Lupinus / Pilosus / Found on y\e / Border of y\e / 
Altamayhaw / J.E” This of course is the Altamaha River, which flows through southeast- 
ern Georgia. The first of these locations is about 360 km northwest of Fraser’s base in 
Charleston; the second is about 240 km to the south. The distance between these loca- 
tions, together with the certainty that these summer-flowering species must both have 
been collected in 1787, isa measure of John Fraser's energy and dedication. 

Most of the specimens (624, or 91%) are labeled with an identification—a genus and 
species, or a genus only. 328 (48%) of the specimens are fully identified (to genus and 
species), by either Walter or Fraser or by both (where Walter wrote the genus and Fraser 
added the epithet). Nearly half of the specimens (296, or 43%) are identified only to ge- 
nus, 183 of them (27%) by Walter, 113 (16%) by Fraser. 64 (9%) of the specimens bear la- 
bels, but were not identified. 


(ea 


ped 
~ 


an the 
identification. Walter frequently appeared not to recognize the plant, but wrote a brief 
Latin diagnosis which reflects his familiarity with Linnaeus’ sexual system of classifica- 
tion. Some examples: 4-C, Ptilimnium capillaceum—“Ammi majus sp. nova?”, 55-C, 
Desmodium cuspidatum-“Hedysarum Flore magnus”, 60-E Silene stellata—“ignota”:; 61b- 
B, Iva imbricata-“lva ? nova”; 67-D, Crotalaria rotundifolia““Lupinus affinis”; 75-F. 
Nelumbo lutea-“The Great Nymphaea’; 86-B, Myriophyllum pinnatum-“Potamogeton 
monoicum”; 87-D, Physostegia virginiana-“Prasium nov”, 89-A, Cynanchum 
angusttifolium—“PentandIria] Digynlial bifollic”; 90-B, Quercus michauxii-“Quercus nova’: 
ae Brunnichia ovat d-“Rajania (monoica)” 97-C, Bumelia lanuginosa-“fol. angustis 
”98-D, Smilax laurifolia-“Smilax Baccis albidis”, 106- 
E, Phyla nodiflo ra-“Verbena nodiflora Didynamia gymnosperm disperma’”, 108-D, Viola 
villosa-“Viola Fol. pubescent”; 114-E, Juncus effusus-“an Juncus”; 117-A, Decodon 
verticillatus-“Decand monogyn nov G. aquatic”; 117-B, Cynoctonum sessilifolium—“Genus 
nov Pentand digyn 

Fraser’s distinctive script usually appears to be of observations made of the plants in 
the field. When he ventures into Latin he is clearly in unfamiliar country. Britten (1921) 
gently noted Fraser's limited linguistic skill where “nova genera” was used “asa singular,” 
trusting his classic-trained readers to understand. Some of Fraser’s notations: 1-D, Vibur- 
num dentatum-‘not in y\e Flora”, 11-B, Asclepias sp.-“Dove Coloured”; 12-G, Aster 
concolor-“Aster Dark Blue Flower”, 22-C, Psoralea canescens—“Buchnera americana na- 
tive name Buck root”, 38-G, Croton punctatus-“Croton Maritimum Nova”, 52-1, (unidenti- 
fied)-“Granadilla not in y\e flora”, 62a-A, Aesculus parviflora—‘Juglans Alba nova”; 67- 
A, Lupinus villosus-“Blue Flowering Lupinus Pilosus Found on y\e Border of y\e 
Altamayhaw J.F”; 67-C, Lupinus perennis~“Lupinus Perrenis [sic]; 72-A, Crataegus crus- 
galli-“Mespilus Large Cock Spur”; 72-B, Crataegus uniflora-“Mespilus Small Cock Spur’; 


The labels of occasional specimens carry comments or information other t 


WARD 1117 


92-K, Rhododendron minus-“Leaves like y\e Kalmia? Latifolia grows on y\e Mountain in 
decid. y\e Est\n & Wes\n Waters”, 93-C, Salix humilis-“Salix Minor? of Fraser”, 100-K, 
Arenaria uniflora-“No Name”; 100-F, Verbena tenuisecta-“Nova Genera”, 108-C, Viola 
primulifolia-“Viola White Flowers Downy Leaf”, 1L0-A, Vitis labruska-“Vitis? Labruska 
Fox Grape”; 116-B, Eriogonum tomentosum-“Sophoranthus nov genera”; 117-C, 
Lachnanthes caroliniana-“Nova Genera”; 117-E, Carphephorus sp.-“Genera Nova.” 

Comments ona few labels are not in Fraser’s hand nor in Walter’s, and show involve- 
ment of a third person: 8-C, Stylosanthes biflora-“Arachis ground nut”, 12-F, Aster sp.- 
“Aster grows 16 feet high”; 46-C, Conoclinium coelestinum—‘Bright violet blue: said to bea 
specific for the venereal complaint”; 105-C, Vaccinium arboreum-“A Vaccinium 5 miles 
from Cranberry measured 50 feet high”; 115-G, Planera aquatica-“Monoecia Triandria G. 
nova.” 

The handwriting on these last-cited labels, and a few others, poses a puzzling, unre- 
solved question. The content of most suggests they were written at the time of collection. 
Their subject matter is wholly different from the technical comments recorded by Walter. 
And the hand is quite incongruous with that of Fraser sr. The “square r” of some (8-C, 12- 
EF 27-D, 38-J, 46-C, 105-C) would seem to be that of Fraser fil. Yet John Fraser, the son, did 
not accompany his father to the Americas until 1800 (Hogg 1852), more than a decade 
after Fraser's 1786-1788 trip, and eleven years after Walter’s death. Could these notations 
have been made by Fraser fil. on his fatlier S ee after ey were brought to England? Or 
is it possible these specimens (and d,and annotated, by Fraser 
fil. on his 1800 trip to the prcienien: with his father? 

The labeling of one species confirms an involvement of a third person in identifica- 
tion and possibly a fourth in mounting of the specimens. A mint, Trichostema dichotomum, 
is represented by two collections, 61b-D and 103-F Both are labeled with its name. The 
label of 103-F is in Walter’s hand, and is spelled conventionally. But 61b-D is in another 
hand, neither Walter's nor Fraser’s. The style of the capital “T” is so different that the 
mounter of the specimens, intending to arrange them in alphabetical order (as was done 
with 103-F), but seemingly misreading the convoluted script of the initial letter, has placed 
6lb-D between “Iva” and “Illicium.” Further, the writer of that label transcribed its epi- 
thet as “dichotoma,” suggesting one of sufficient education to perceive an apparent error 
of gender mismatch between epithet and genus, yet without adequate classic background 
to understand that “-stema” is a neuter root. Even more, the mounter himself must have 
been of limited experience not to have recognized the distinctive specimen as one al- 
ready found elsewhere in the folio. With certainty, Fraser sr. neither formed this label nor 
was involved in its mounting. 

One specimen is wholly aberrant. Spm. 102-A isa branched structure mounted alone 
ona full page. It is a gorgonian (Gorgoniidae-coral, s.L.). Enlarged bodies on its branches 
are barnacles (Archaeobalanidae; Conopea sp.). It was labeled “Sea Plant’ in Fraser’s hand. 

The physical processing of the specimens into the present folio remains poorly un- 
derstood. Walter, of course, was not involved. Fraser, perhaps soon after his return to En- 
gland in March 1788, must have acquired the empty folio and, as indicated by his writing 
on the title page, established its goal. It is often forgotten that the present herbarium of 
690 plants is only a subset of the “upwards of thirty thousand dried specimens of plants” 
Fraser (1789) claimed to have gathered during his 1786-1788 trip to the Americas. An 
unknown number, perhaps the greater part, were sold by Fraser (1789) to Charles Louis 
LHeritier, a wealthy French botanist, and are now in the Lamarck herbarium, Paris. The 


1118 BRIT.ORG/SIDA 22(2) 


basis is unknown by which Fraser, or his sons during his absence on later trips to the 
Americas, Cuba, and Russia (Hogg 1852; Simpson et al. 1997), selected from this larger 
collection the plants to be retained in the “Walter Herbarium.” 

The historic importance of this early sampling of the plants of Georgia and the Caro- 
linas is unquestioned. America is fortunate the fates have preserved this fragmentary 
glimpse of its vegetation as it appeared before the full impact of modern civilization. 
Even so, the larger value of the John Fraser folio and the specimens it contains lies in the 
degree it supports the writing of Thomas Walter and his Flora Caroliniana. The nomen- 
clatural basis for Walter's work and its relation to Fraser’s herbarium is to be discussed in 
subsequent reports of the Thomas Walter Typification Project. 


ACKNOWLEDGMENTS 


[remember with gratitude the courtesy shown my wife and me in 1984 by N.K.B. Robson, 

Mike Mullin, and other members of the staff of the Natural History Museum, London, 

and especially the salutation given me by William T. Stearn with his pleasure in meeting 

the “Anonymos botanist.” | have been immeasurably aided by the Bernice Schubert pho- 

tos whose negatives were loaned me in 1977 by Carroll E. Wood, Harvard University. I 

have been given assistance in matters of history, linguistics, and taxonomy by Charles E. 

Jarvis, Natural History Museum; Alexander Krings, North Carolina State University; James 

S. Pringle, Royal Botanic Gardens, Hamilton, Ontario; David H. Rembert, University of 

South Carolina; and Robert L. Wilbur, Duke University. 

REFERENCES 

Buake, S.F.1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137. 

Britten, J. 1921. Thomas Walter (1740?-88) and his grass. J. Bot. 59:69-74. 

Coker, W.C. 1910.A visit to the grave of Thomas Walter. J. Elisha Mitchell Sci. Soc. 26:31-42. 

Ewan, J. 1979. Addenda on Thomas Walter (c. 1740-1789). Taxon 28:345-346. 

Faben, R.B. 1989. Commelina caroliniana (Commelinaceae): A misunderstood species in the United 
States is an old introduction from Asia. Taxon 38:43-53. 

FerNatp, M.L.and B.G. ScHuserr. 1948. Studies of American types in British herbaria. Part IV:some spe- 
cies of Thomas Walter. Rhodora 50:190-208, 217-229. 

Fraser, J. 1789. A short history of the Agrostis Cornucopiae: or, the new American grass. Chelsea, 
England. 

Gray, J. L., ed. 1893. Letters of Asa Gray. Boston. 

HiicHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rep. 16:31-56. 

Hoc, R. 1852. Life of John Fraser. Cottage Gardener 8:250-252. 

Maxon, W.R. 1936. Thomas Walter, Botanist. Smithsonian Misc. Collect. 95:1-6. 

RemBert, D.H. 1980. Thomas Walter, Carolina botanist. South Carolina Mus. Comm., Bull. No. 5:1-33. 

Simpson, M.B., S. Moran, and S.W. Simpson. 1997. Biographical notes on John Fraser (1750-1811): plant 
nurseryman, explorer, and royal botanical collector to the Czar of Russia. Arch. Nat. Hist.24:1-18. 

Smait, J.K. 1935. Thomas Walter's botanical garden. J. New York Bot. Gard. 36:166-167. 

Walter, T. 1788. Flora Caroliniana. London. 

Warb, D.B. 1962. The first record of the Fraser Fir. Castanea 27:78-79. 

Ward, D.B. 1977. Nelumbo lutea, the correct name for the American lotus. Taxon 26:227-234. 


— 


EVIDENCE FOR HYBRIDIZATION BETWEEN TWO SYMPATRIC 
VIOLET SPECIES, VIOLA GRAHAMII AND V. HOOKERIANA 
(VIOLACEAE), INCENTRAL MEXICO 


Aurea C. Cortés-Palomec'! Harvey E. Ballard, Jr. 
Department of Environmental and Plant Biology Department of Environmental and Plant Biology 
i iversity hio University 
Athens, Ohio 45701, U.S.A. Athens, Ohio 45701, U.S.A 
ac399295@ohio.edu ballardh@ohio.edu 
ABSTRACT 


Viola hookeriana and V. grahamii (Violaceae) are closely related species in section Viola, subsection Mexicanae 
occurring in POENTAO Ue habitats across much of central Mexico. In various locations, these species grow sym 
many individuals with intermediate morphological enasctes has ee 


jah 


eee and 
een between these species. Using a combination of nao POE ecological, and Oeulg evidence 


we evaluated two pmined populalions of V. ce ahamiiand V.} Lake Patzcuaro 
in the state of N I tential of detec hiv bud morphologies, the strength of phenologi- 


cal differences and ecological Re aen among the species and putative hybrid, and to Saini the level of 
gene flow between the two species. Our results indicate that hybridization is equ and that hybrids are 


morphologically distinct from the parental taxa. Pre-mating isolation is weak and has favored extensive hybrid- 


ization due to largely overlapping blooming times, weak ecological isolation and absence of spatial isolation. 


Hybrids are morphologically and genetically closer to Viola grahamii, but do show unique alleles as well. 
yy, Pp g y g y 8 q 


RESUMEN 


AA rr J | 


Viola hooke iy Vgral ii (Viol ) | pecies f la Subseccién 
la Seccion Viola del género Viola. E | | México 


y la presencia de individuos con eee nent ee ha sugerido la existencia de hibridizacion entre las 
especies. Usando una combinacion de datos morfolégicos, ecolégicos y moleculares evaluamos dos eee 
mixtas de V.grahamii y V. hookeriana en i. montanias cercanas al lago de Patzcuaro en el estado de Michoacan 
en México para determinar la presencia de morfologias caracteristicas a los hibridos, el grado de diferencias 
fenoldgicas y ecologicas entre las especies y el nivel de flujo génico entre ellas. Nuestros resultados indican la 


presencia de hibridos y que estos son mor sol Opicamente sae a bi boos Los mecani le aislamiento 
+ 1] ]} 


entre las especies parenta tales noson muy f fuertes y 
tiempos de floracién similares, muy poc ient logico y la ausencia de separacion espacial entre las 
especies. Los hibridos son eee y genéticamente mas similares a Viola grahamii, pero muestran 


aleleos que son unicos a ellos. 


Hybridization isa common phenomenon in nature, as evidenced by an estimated 70,000 
natural interspecific plant hybrids worldwide (Rieseberg & Ellstrand 1993; Judd et al. 
1999), and the fact that between 16 and 37% of the plant families reported in different 
floras contain at least one hybrid taxon (Ellstrand et al. 1996). In some cases, hybridiza- 
tion can lead to the creation of hybrid species and/or intr ogressants between hybrid de- 
rivatives and the parental taxa (Arnold 1992; Rieseberg 1995). The genus Viola L. 
(Violaceae), which comprises about 525-600 species (Clausen 1964; Ballard et al. 1999), is 
well known for its taxonomic problems due to hybridization and introgression, as well as 
complex patterns of variation in individual traits (Brainerd 1924, Anderson 1954; Rus- 


'Current Address: C Ecosistemas, Uni idad Naci | tc Je Méxic , Antigua Carretera a 
Patzcuaro No. 8701, Col. Ex-Hacienda de San José de la Huerta, Morelia, 58190 Michoacan, MEXICO, 


SIDA 22(2): 1119- 1133. 2006 


1120 BRIT.ORG/SIDA 22(2) 


sell 1955; Ballard 1994; Krahulcova et al. 1996; Gil-ad 1998; Neuffer et al. 1999; Marcussen 
& Borgen 2000; Calderon de Rzedowski 2001; Harmaja 2003). 

Analysis of herbarium specimens from central Mexico has indicated that hybrid- 
ization is likely occurring among species belonging to Viola subsection Mexicanae (Bal- 
lard 1994). Viola subsection Mexicanae W. Becker (sensu Ballard et al. 1999) is a mono- 
phyletic group consisting of eight stoloniferous or rosette-forming species: Viola 
hemsleyana G. Calderon, V. grahamii Benth., V.oxyodontis H.E. Ballard, V. humilis H.B.& 
K., V. hookeriana H.B.& K,, V. beamanii G. Calderon, V.guatemalensis W. Becker, V.nannei 
Polak. (Ballard & Sytsma 2000; Ballard et al. 1999), and an additional undescribed spe- 
cies (Ballard, per. obs.). Instances of hybridization have been identified from several lo- 
calities in central Mexico occurring between Viola grahamii and four other members of 
the group, V.guatemalensis, V. hemsleyana, V. hookeriana and V. humilis. Occurrences of 
these putative hybrids appear to be restricted however to the immediate areas where both 
parents occur sympatrically. 

Viola grahamti (known in Mexico as “hoja de pasmo” or “pensamiento del cerro”) is a 
frequent to locally abundant perennial distributed across much of Mexico. Specifically, 
it occurs in mountainous regions from northern Mexico south to northern Guatemala, 
and is one of the most widely distributed members of the subsection Mexicanae (Fig. 1). 
It thrives in dry to mesic sandy loam under varied forest canopy and along stream banks 
at elevations of 1950-3600 meters (Ballard 1994; Calder6n de Rzedowski 2001), and often 
forms large mats connected via aboveground stolons. It produces chasmogamous flow- 
ers at the beginning of the rainy season June) for approximately four weeks and cleisto- 
gamous flowers for several months afterward June-December) until the onset of the dry 
season (Cortés-Palomec 2005). 

As previously mentioned, one of the four species with which Viola grahamii appears 
to hybridize is V. hookeriana (known in Mexico as “violeta”). Viola hookeriana is an infre- 
quent to locally common perennial generally growing in small isolated populations across 
northern and central Mexico (Fig. 1). It grows in mesic loam under mixed deciduous and 
coniferous forest canopy at elevations between a 2500 meters oe 1994; Calderon 
de Rzedowski 2001). Reproduction occurs via both ¢ d clei flow- 
ers and follows a similar pattern to that seen in V.grahamii (Cortés- Palomec, pers. obs.) 

Viola hookeriana and V. grahamii are divergent in several characters: growth habit, 
flower structure, and foliage, and can be separated without difficulty in both herbarium 
specimens and living populations (Table 1) (Ballard 1994). The putative hybrids have been 
identified in areas of overlapping distribution of the two species (Fig. 1), and are recog- 
nizable as exhibiting intermediate morphologies (Ballard 1994). 

The goal of our study was to assess the presence and extent of hybridization between 
V. grahamii and V. hookeriana in natural populations, and identify intrinsic pre-mating 
isolation mechanisms that may be functioning between the two species. We examined 
morphological and phenological differences between the two species, characterized the 
ecological factors governing where individual species and putative hybrids occur, and sur- 
veyed genetic patterns to identify gene flow between species in two study sites in central 
Mexico where V. grahamii and V. hookeriana intermingle extensively. 


MATERIALS AND METHODS 


Site Establishment and Sampling 
Two study sites were established in the mountains to the north of Lake Patzcuaro in the 


CORTES-PALOMEC AND BALLARD 1121 


A Viola grahamii 
A Viola hookeriana 
© V. grahamii x hookeriana 


Study sites Su 
Northern Michoacan os 


0 500 Km 


Fic. 1. Distribution of Viola grahamii, V. hookeriana, and V. grahamii < hookeri Mexi 1 I la. Distribution 
from Ballard (unpublished data). 


municipality of Quiroga, Michoacan, Mexico (Fig. 1). The sites were established in early 
June 2000 and observations were made weekly over a period of five weeks during the 
summer rainy season when chasmogamous flower production was most extensive. The 
two specific sites were chosen due to the presence of abundant populations of the two 
species and their putative hybrids as identified through a survey of the area conducted 
earlier in the spring. The first site (Site A) [19°41'57" N; 101°32'27" W] was situated on the 
lower southern slopes of Mt. Zirate, above the town of Santa Fe de la Laguna. The second 
site (Site B) [19°42' N; 101°35' W] was located 8 km to the west of Santa Fe de la Laguna 
(Km 48 on the road from Quiroga to Zacapu (MX-15)). While both sites were located in 
superficially similar environments, they differed in the level of disturbance and abun- 
dance of violet species. Viola grahamii was abundant in both sites, but areas of visually 
pure V. hookeriana separated from V. grahamii were found only in Site A. 

In Site A, four 20 m long transects were established. Two transects were arbi 
through the middle of a visually “pure” population of the parental species (A-T1VG ina V. 
grahamii population and A-T4VH ina V. hookeriana population). Two additional transects 
(A-T2HY and A-T3HY) were placed through areas where putative hybrid plants inter- 
mingled with the parental taxa. Ten quadrats (0.5 m2) were randomly established along 
each transect with position determined using a random number table. In Site B the same 
procedure was followed except that only three transects were positioned resulting in a to- 
tal of 30 quadrats (B-T1VG for V. grahamii, B-T3HY of mostly V. hookeriana and a mixture 
of V.grahamii and hybrids, and B-T2HY consisting mostly of putative hybrids). 


ily placed 


1122 BRIT.ORG/SIDA 22(2) 


Taste 1.Comparison of morphological characters used to distinguish between Viola grahamii and V.hookeriana 
(on the basis of Ballard 1994). The characters of hybrids are not included since they are not consistent (i.e, 
hybrids show varying combinations of traits, making them difficult to characterize). 


Character Viola grahamii Viola hookeriana 

Stolon position above ground underground 

Stolon nodes present absent 

Stipule adnation free or adnate free 

Pubescence on petioles and present [abundant] absent 

abaxial surface of leaves 

Leaf shape ovate-oblong to elliptic broadly ovate to reniform 
Length of pedicels (cm) 3-12 5-21 

Calyx pubesence ciliate glabrous 

Corolla color white white, sometimes purplish 
Elevation 1950-3600 m 1700-2500 m 


For transects representing “pure” parental taxa at each study site, three plants that 
were not connected via stolons were randomly selected in each of the 10 quadrats. One 
leaf was removed from each of the three plants and dried in silica gel for later DNA ex- 
traction. A second leaf, the largest of each individual, was collected and labeled to corre- 
spond to the DNA sample. This leaf was pressed and used for subsequent morphological 
analysis. The same methodology was used for plants in the putative hybrid/mixed-taxon 
transects, except that 5 individuals were sampled per quadrat. 


Phenology and Pollinator Visitation 

In each of the 70 quadrats (40 from Site A, 30 from Site B), the number of individuals 
present per quadrat and the number of individuals with open chasmogamous flowers 
were recorded weekly during the five weeks of field observations to interpret phenology. 
The percentage of flowering individuals, those bearing chasmogamous flowers at some 
point during the study, was calculated for each visit for the two species and for the puta- 
tive hybrids to allow for a comparison of phenology among the parental taxa and puta- 
tive hybrids at both sites. ae with more than one taxon were recorded separately 
for each taxon and analyzed as te observations. A Kolmogorov-Smirnov two-sample 
test was performed in NCSS (Hintze 1999) to compare phenological patterns among taxa 
at each of the two sites. 

Pollinator visitation was assessed during weekly visits to Site A, which was the only 
site to have clearly differentiated populations of V.grahamii and V. hookeriana. A total of 
40 hours of direct observations of the flowering individuals were recorded with special 
emphasis made to observe instances of pollinator movement between or among differ- 
ing taxa. The hours of pollinator observation were equally divided between two daily 
periods relating to potential insect activity based on the times when previous observa- 
tions in the area suggested the potential pollinators would be more active, between 8:00 
and 11:00 am, and between 12:00 and 3:00 pm. 


as 


Ecological Characterization 

To determine if the taxa grow in modally different microhabitats, infer whether ecologi- 
cal isolation might reduce gene flow between the species and the putative hybrids, and 
understand the local distribution of putative hybrids relative to the parents, an ecologi- 
cal characterization of microhabitat in the 70 quadrats was conducted. Soil N, P, K con- 


CORTES-PALOMEC AND BALLARD 1123 


centrations and pH were evaluated using a LaMotte™ Combination Soil Test Kit (LaMotte, 
Chestertown, MD). Values were given in exact units for pH while values for N, P and K 
were determined via a colorimetric test procedure from which concentration could be 
estimated within a set range, 5-75 ppm for N, 5-100 ppm for P, and 50-200 ppm for K. Due 
to the inability of obtaining exact concentration values for N, P, and K, these values were 
range standardized and analyzed in rank form. 

Percent soil moisture was recorded using the gravimetric method described by Hadley 
and Levin (1967), with the modification that soil mass was determined after one week of 
air drying. Light availability as a direct means of inferring canopy closure was measured 
using a Model-C spherical densiometer (Vora 1988). Quadrats with more than one taxon 
were recorded separately for each taxon and analyzed as separate observations. Data were 
standardized and an analysis of variance was performed in NCSS to test for differences 
among V. hookeriana, V.grahamii and putative hybrids. 


Morphological Analysis 

From the pressed leaves, measurements were taken of the 0°, 30°, 60°, 90°, 120°, 150° and 
180° radii, following the procedure outlined by Ballard and Wujek (1994). The center of 
the leaf was considered to be the point along the midrib opposite the widest part of the 
leaf blade. Presence or absence of leaf pubescence on the abaxial surface of the leaf, a 
characteristic considered diagnostic for separating V. hookeriana (glabrous) and V. 
grahamii (pubescent), was also recorded. Measurements were standardized and a dis- 
criminant analysis (Manly 1994) was performed, both including and excluding visually 
identified putative hybrids to determine overlap in leaf characters among the two taxa 
and putative hybrids. The analysis was performed in NCSS using an automatic variable 
selection procedure. Measurements at 30° and 120° were excluded from the analysis due 
to colinearity of the data. The canonical scores generated from these analyses were used 
in an analysis of variance (ANOVA) with a Fisher’s LSD pairwise comparison analysis 
(Zar 1996). 


Genetic Analysis 

To test the presence of genetic differences among the parental taxa and identify potential 
gene flow within the putative hybrids, biparentally inherited nuclear markers, inter- 
simple sequence repeats (ISSRs) were used. ISSR data have proven to be highly effective 
in detecting hybridization and/or gene flow among closely related species, and to test 
the hybrid speciation hypothesis (Wolfe et al. 1998a, b, Wolfe & Randle 2001; Archibald 
et al. 2004; James & Abbott 2005). Additionally these markers are useful in studies of 
differentiation among closely related species for they are able to utilize some of the vari- 
ability present at microsatellite loci without the need to develop species-specific 
microsatellite primers, an advantage when working with species which have not been 
well-studied genetically. 

Genomic DNA was extracted from the silica gel-dried leaves using a Wizard Genomic 
DNA purification kit (Promega, Madison, WD). Five different ISSR primers (McCauley & 
Ballard 2002) were initially screened to identify primers which would produce consis- 
tent and scorable polymorphic bands among the taxa. Two primers were selected: Wolfe 
#99B [(CA)6GG] and HB#15 [(GTG)3GC]. These were used for amplification of ISSR prod- 
ucts in replicated 25 ul reactions consisting of 1 wg wl! diluted genomic DNA, 19 pl auto- 
claved distilled water, 2.4 n110x PCR Buffer (Gibco BRL), 2 41 MgCl2 (50 mM, Gibco BRL), 
2 ul of dNTP mix (10 mM, Fisher), 0.5 ul of BSA (Bovine Serum Albumin, 4 ug pl Fisher), 


1124 BRIT.ORG/SIDA 22(2) 


0.25 wl primer and 0.25 wl of Taq polymerase (5 U 11, Gibco BRL). The polymerase chain 
reaction was performed in a Stratagene RoboCycler 96 with hot-top (Stratagene Inc, La 
Jolla, CA) and programmed for 2 min. at 94°C; 40 (primer Wolfe #99b) 44 (primer HB 
#15) X 30 sec. at 94°C, 45 sec. at 44°C, 1 min. 30 sec. at 72°C; 20 min at 72°C. 

PCR products were electrophoresed in a 1.3% agarose gel in 0.5 x TBE buffer with 
flanking 250 bp ladders (Gibco BRL). Gels were stained with a solution of ethidium bro- 
mide in 0.5 x TBE buffer for 20 minutes and imaged under UV light. Gel images were 
analyzed with BioMax ID image analysis software (Version 2.0.3, Eastman Kodak Com- 
pany, Rochester, NY) to identify and size fragments. Fragments were scored as present (1) 
or absent (0), with fragments comigrating at identical rates (+ 10 base pairs, the general 
limit of resolution for agarose gels) considered equivalent. Statistics concerning fragment 
occurrence, including total fragment number, distribution across taxa, polymorphic and 
fixed fragments, and number of fragments shared among taxa per site were calculated 
using TFPGA 1.3 (Miller 1997). 

Analysis used three discrete methods to evaluate relationships and similarity among 
the taxa and putative hybrids. UPGMA cluster analysis and Principal Coordinates Analy- 
ses (PCoA) were performed both within and between the two sites using NTSYS ver. 2.02} 
(Applied Biostatistics Inc.). An Analysis of Molecular Variance (AMOVA) was addition- 
ally performed ina hierarchical fashion among species and sites using WINAMOVA 1.55 
(Excoffier 1993). All analyses were performed according to the methods described in 
McCauley and Ballard (2002). 


RESULTS 


Phenology and Pollinator Visitation 

The parental spec d divergent but non-significant tendencies in blooming time; 
in fact, in Site A V. hookeriana bloomed first, whereas in Site B it bloomed later (Fig. 2). 
The Kolmogorov-Smirnov two-sample test showed no significant differences among the 
blooming times of the hybrids and the parental taxa in either site (p values = 1 in some 
pairwise comparisons) (Fig. 2). No pollinators were observed visiting violets at any time 
during this study. 


Ecology 

All three taxa inhabited modally different microhabitat conditions influenced by light 
and certain nutrients, in all cases where there were significant differences between the 
taxa, the hybrids grew in intermediate conditions to the parental species. The ANOVA of 
the ecological factors revealed significant differences in light (p= 0.02), P (p = 0.04) and K 
(p = 0.03) among V. hookeriana, V.grahamii and the putative hybrids; moisture (p= 0.32), 
pH (p=0.86) and N (p= 0.62) did not differ significantly. Viola hookeriana grew in higher 
concentrations of P, lower concentrations of K and more shaded environments than V. 
grahamii (Table 2). 


Leaf Morphology 

Two hundred and eighty eight individuals were classified into three groups correspond- 
ing to the two parental taxa and the putative hybrid individuals (Fig. 3). The discrimi- 
nant analysis indicated three marginally distinct groups (Fig. 4), with the GLM ANOVA 
and the Fisher’s LSD Multiple Comparison analysis indicating that they were well-sup- 
ported and significant (p < 0.05) groups. 


CORTES-PALOMEC AND BALLARD 


% of flowering individuals 


% of flowering individuals 


70 
Site A V. hookeriana AT4 
KZA hybrids AT2 
60 + hybrids AT3 
SS V. grahami AT1 
“| i. AT1 AT2. _AT3 
me AT2 77 
AT3 1 1 
AT4 77 22 HE 
40 + 
SO" 
20 + 
i : ] 
0 md 
4 July 6 July 12 
Visits 
70 . 
Site B (—] V hookeriana BT1 
V-ZiZA hybids BT2 
60 4 RSs9_:V grahami BT3 
BT1._ BT2 
50 - BT2 .873 
BT3 1.873 
40 4 
30 4 
20:4] 
1c 
0 ; iA 


June 25 


! 
July 12 July 18 


Fic. 2. Histogram of phenology for fl 


1125 


J 


t 
Vu Vb L 


als sampled are as f 


rs 
JTHITTOY 


ollows (num 


twU 


ber of reproductive individu 
VG(AT4)=146(77). Site B VG=35(19); HY=95(55); VH =142 (33). 


Ve. FIUURCHIGH 


als): Site A: VH(AT 


jana, VG=V.g 
1)=24(12); HY(AT2)=151(76); HY(AT3)=232(134); 


1126 BRIT.ORG/SIDA 22(2 


Taste 2.Means and SE for < Se. lected eG ologic al parameters aMONng V. hookeriana (VI 1), \ ahamii (\ G) a and h ybr ids 
(HY). Superscript letters indicate a significant difference for a particular parameter among the taxa. Moisture 
light are expressed as percents. P N and K values are along a scale of | to 7 (l=very i 7= high). 


anc 


Taxon Moisture Light pH P N K 
VH 26(+£0.1) 16(£0.4)" 6.40(+0.14) 5.32(£0.49) 1.28(+0.10) 6.44(+0.16) 
VG 24(+1.1) 29(£0,3)’ 6.51(#0.12) 3.82(+0.37)° 1. 


14(+£0.09) 7, O(+0.13)° 
HY 25(+0.1) 29(£0.3) 6.43(+0.10) 4.40(+0.33) 1.18(+0.08) 6.81(+0.12) 


Genetic Data 

A total of 242 Viola individuals were examined genetically with ISSRs: 33 Viola 
hookeriana in Site A and 19 in Site B, 65 V. grahamii individuals in Site A and 35 in Site B, 
and +8 putative hybrid individuals in Site A and 42 in Site B. In Site A, 70 ISSR fragments 
were resolved vs. 53 in Site B. The majority of these were identical between the two sites, 
resulting in a total of 75 unique and scorable fragments for the study sites together. The 
relative frequency of individual fragments ranged from less than 10% of individuals (rare 
to infrequent) to 100% (ubiquitous). Distribution of fragments across taxa (Table 3) showed 
that some were specific to one of the parental taxa (e.g. five fragments were species-spe- 
cific for V.grahamii,and three were specific for V. hookeriana) whereas others were shared 
among taxa. Most of the shared fragments occurred between V. grahamii and the puta- 
tive hybrids (e.g., 17 fragments in Site B). Nine fragments were specific to the putative 
hybrids. 

UPGMA cluster analysis (Fig. 5) showed strong separation of the two study sites. All 
individuals diverged substantially between Site A and Site B, regardless of taxon. The 
PCoA confirmed these discrete site differences (Fig. 6A). Additional PCoA analyses were 
carried out separately on the two sites [Site A (Fig. 6B) and Site B (Fig. 6C)]. In both sites, V. 
grahamii and the putative hybrids showed the most extensive genetic overlap. In Site A, 
V. hookeridana mostly segregated from the other taxa, although some individuals clus- 
tered with putative hybrid derivatives, and others with V. grahamii. In Site B (Fig. 6C) 
two distinct groups of V. hookeriana were evident, with some putative hybrids occurring 
within each of them. In all cases, putative hybrid individuals were widely placed and 
substantially overlapping in distribution with the parental taxa, together expressing a 
greater level of genetic diversity. 

The AMOVA (Table +) showed that most of the variation was within morphologi- 
cally defined taxa (69%), although significant differences were detected between sites 
and among taxa. Variation between sites accounted for 27% of the total variance. Differ- 
ences among parental taxa and hybrids accounted for only 3.72% of the variance. The 
same pattern was observed when the two sites were analyzed separately. In Site A, 94.5% 
of the variance was due to variation within the taxa; 95.3% in Site B. If the hybrids were 
eliminated, variation was slightly reduced within taxa (90.32% and 89.17%); however, lev- 
els of variation within the species themselves remained high. 

DISCUSSION 
Our results show conclusively that hybridization is occurring between V. grahamii and 
V. hookeriana. Phenetic analysis of leaf shape characters in the species, more exploratory 


than exhaustive, showed separation among V. hookeriana, V. grahamii and the hybrids. 
Hybrid individuals formed a distinct group from the parental taxa suggesting that leaf 


1127 


CORTES-PALOMEC AND BALLARD 


iu hookeriana, 


b 


Fic. 3. Rey 


Axis 1 


PET] Oa ee Pe en Pa Pee 
\ i‘ 


a) 
s 
4 
-t 
“ 
~ 
al 
—d 
So 
sc 
Pa 
oO 
pes 
L=3) 
= 
S 
= 
= 
xg 
cs YY 
a 
ba) 
=) 
ae 
= 
<x é 
<< S 
3 8 
a DD 


1128 BRIT.ORG/SIDA 22(2) 


Taste 3. Number of ISSR bands found per site per taxon (V hookeriana, V. grahamii and hybrids), as well as num 
ai of bands shared among the taxa. 


Number of bands SITEA SITEB 


atl 


Viola hookeriana 3 

Viola grahamii 5 5 
hybrids 3 

V. hookeriana and hybrids | 

V.grahamii and hybrids 11 7 
All individuals 46 20 


VHB 
| q T T ! T T qT qT | T T T T 1 T T rr a | 
-0.01 0.09 0.20 0.30 0.40 
Genetic distance 
Fic. 5, UPG d tionshi g taxa a transects. Similarity compared using the AMOVA-derived PhiST 
distance matrix. (VHA = Viola hookeriana in Site A, HYA = hybrid : V. grahamii in Site A; tt f for Site B) 


traits are useful in distinguishing the hybrids from the pure species, similar to the find- 
ings seen in other Viola (Russell 1954; Jonsell et al. 2000). While these differences appear 
to follow a clear pattern, our potential discovery of introgressant individuals may actu- 
ally blur the distinction to some degree. Extensive gene flow was confirmed, and has 
proceeded to the extent that the two sites harbored substantially different genotypic com- 
binations. At both sites, numerous morphologically “pure” parental individuals of the 
two species were demonstrated to be cryptic hybrid derivatives, with a preponderance 
of these resembling V. grahamii. High levels of genetic variability within taxa may be the 
product of long-continued gene flow. The results from AMOVA suggest that around 95% 
of the genetic variation is due to variability within each taxon. While this high level of 
variation could be partly artifactual, owing to rapid evolution of ISSR primer sites, it is 


es 


CORTES-PALOMEC AND BALLARD 


1129 


06 
04 4 age 
oe 
(o) 
eC%y o ® 0 2D o 
* e og @ 
=a 024 ° ee, °9 coor, ® 
xo (o) 
2 @ Po Qo Pel ayo 
) fo) fe) 00 Oa 
~ 004 % % o a Bee op 
N oe ra) ‘e) 
® ° Bae 8 ° oo 08 6 
@ O fo) ees 
% oo] 2 Op @ OW Oraree tg 
cate) a) e 
2) e Oo 
oO m0 Ce &@ 
aee. ° 
0.4 4 g 
-0.6 : ' : : 
-0.6 0.4 -0.2 0.0 02 0.4 06 0.8 
Axis 1 (12.18%) 
06 
B e 
e sd 
044 e 
a ee %, e°, 
° ¢ 0 
oO e e* 
024 — mom ° "5 6 ® 
x fe) e° * S 2) Q O ie) 
6 De SoM awe 080 9 oo Ba 
= 00 4 e oBo o oo 
N 0 Sg @ 00 pg ° 0 
2 & 60 9 © 
< i fo) 8, °% ° 
foie) fe) 
0.2 4 oo F ° © »6%0o 
SY 2 6 
‘ fo) 
08 5 fe) 
044 9 oa) 
° fe) 
-0.6 T 1 T T T 
-0.6 -0.4 0.2 00 0.2 0.4 0.6 
Axis 1 (7%) 
0.6 
C 0 
04 4 ° fo) 
(e) ® ; s 
Po o © eo 
0.2 4 2 o # fe) °° 
S : ° ° @ fo) 
ae % ° 
re) ray o) Q © (3) O° 
pe e@o0 o 0 oe 
~ 004 °., 3 fe) 
N oe oO 1a] 
g fo) e 6 * e of 2 
ed % 2 : 
-0.2 4 Oo O 
os eo 
ef 0G @ 
fo) e 
-0.4 4 ® * 
-0.6 T T T T 7 
-0.6 04 0.2 0.0 02 04 06 
Axis 1 (10%) 
Fic . iva . +. A I} . {DfAA\ £ - J * +b. 1 J 4 es | res £, 4 J | Wl 7 | j eee ey 
sites (A and B). B) PCoA of individuals collected in Site A. C) PCoA of individuals collected in Site B. P tag h axis indicat 
- £ ting! ‘elie tg pl > 1 by tk rs = trea g L bh ie re ep os, ee Pe! L Jk 1 J \ 1V.g LL ii x hookeriana 


r J 


(gray dots). 


1130 BRIT.ORG/SIDA 22(2) 


Fase 4. Analysis of Molecular Variance (AMOVA) of 242 individuals collected from both sites (A and B), based on 
2000 permutations. Mean squares (MDS), variance component and percentage of variance as well as P values 
are presented. A nested design was performed. 


Source of variation df MDS Variance component Percentage p 

Between sites | 286.41 2.30 26.79 < 0.0005 

Among taxa 4 18.38 0.32 Se < 0.0005 

Within taxa 236 5.98 5.98 69.49 < 0.0005 
24) 


more likely that hybridization and subsequent gene flow have proceeded to the point 
that many morphologically “pure” parental individuals are actually products of extensive 
gene flow, bridging the gap between parents and hybrids and obscuring taxon boundaries. 

Additional genetic evidence suggesting that the hybridization between V grahamii 
and V. hookeriana has likely been occurring over a long period of time is the presence of 
ISSR fragments unique to the hybrids. It has been suggested by various authors that later- 
generation backcrosses might contain only a small number of the fragments from the 
parental species (O'Hanlon et al. 1999). If this is the case, the development of fragments 
that are “specific” to the hybrids are best interpreted as the result of later generation re- 
combination and long-term post-hybridization genetic reorganization, as suggested by 
O'Hanlon et al. (1999). Studies of introgression in German violets conducted by Neuffer 
et al. (1999) using RAPDs describe a similar phenomenon in the V. riviniana Rchb—V. 
reichenbachiana Jord. ex Bor. complex, but the fact that no unique hybrid bands were 
found led them to conclude that the hybrid complex hada recent origin. Directional back- 
crossing has also been suggested towards V. lutea subsp. sudetica (Willd.) Nyman in hy- 
brids of this species and V. tricolor L (Krahulcova et al. 1996). In this study, the presence 
of many “hybrid-specific” bands suggested a significant time component in the forma- 
tion of hybrid derivatives as well as differentiation in some. Our results suggest that in- 
trogression is occurring between V. grahamii and V. hookeriana, and that hybridization 
might have taken place thousands of years in the past or may occur only rarely, but high 
evels of interfertility and appropriate site conditions have allowed the hybrids sufficient 
time to backcross with the parental taxa. 

Hybridization has likely been favored by the absence of a temporal reproductive iso- 
lation between the two species. Despite a minor, non significant, shift in blooming times 
of the two species (between populations) they have greatly overlapping blooming times, 
which provide extended opportunities for inter-taxon crossing. This small shift in bloom- 
ing may be related to differences in microsite characteristics and levels of human pertur- 
bation such as wood cutting for local uses, which were more pronounced in Site B than in 
Site A. The differences in canopy openness between the two sites due to human distur- 
bance can also explain why V. hookeriana was not common in Site B, while V. grahamii 
was very abundant. Open environments with more light favor V. grahamii, which grows 
best in open-canopy forest or open sites while V. hookeriana is most abundant in more 
shaded environments, mainly under the forest canopy. Other differences were found in 
terms of soil requirements; V. hookeria nd SrOWs in higher concentrations of Pp and lower 
concentrations of K than does V. grahamii. Soil nutrients in the area however are highly 
heterogenous (Cortés-Palomec 2005), and while they may be affecting the distribution 


—" 


— 


CORTES-PALOMEC AND BALLARD 1131 


of V. hookeriana and V. grahamii, the hybrids appear to survive in various soil condi- 
tions, accounting for their wider local distribution in each site, especially in areas with 
moderate to heavy levels of disturbance. It is commonly suggested that since hybrid in- 
dividuals have partial genotypes from both parental species they can potentially grow in 
both parental habitats as well as in their own “intermediate” niche and even lead to the 
extinction of the parental taxa if one of them is rare (O'Hanlon et al. 1999; Levin et al. 
1996). In fact, hybrids between V.tricolorand V. lutea subsp. sudetica have been suggested 
to have a wider range of distribution than both parental species (Krahaulcova et al. 1996). 
In addition, disturbance can lead to “hybridization of the habitat” (Anderson 1954; 
Rieseberg & Ellstrand 1993), resulting in a mosaic of microhabitat conditions that favor 
hybrids and in extreme cases may cause the local extinction of one or both of the paren- 
tal taxa (Rieseberg & Ellstrand 1993; Rhyme & Simberloff 1996). Hybrids in this case can 
be compared to alien plants in the sense that they are new elements in the flora that can 
threaten the original flora of the region (Neufer et al. 1999). Perturbation was high in 
both study sites, mainly due to human activity (ie. livestock grazing and timber har- 
vest). If we consider our sampling of the study sites as representative for the region, then 
the hybrids may be said to be as abundant or even more abundant locally in the zone of 
sympatry than the parental taxa. The heavy genetic overlap of hybrids with V.grahamii 
may threaten that species with local genetic swamping, but since it is widely distributed 
and occurs as pure populations across large regions of Mexico and northern Guatemala 
that is probably not a problem. Given the large number of hybrid individuals between 
the two species at the sites, and the comparatively small number of V. hookeriana plants 
at any given site, however, the latter species may indeed become imperiled (Levin et al. 
1996) 

Sharing of similar flowering times has likely favored pollen movement between the 
two species, however pollinator activity would play a key role in successfully moving the 
pollen. During the year this study took place, several potential pollinators were seen in 
the area of study, however none of them were observed visiting Viola, a finding which 
seems to be abnormal. Similar observations on blooming individuals of one of the spe- 
cies (V. grahamii) over the next two subsequent summers did however reveal the pres- 
ence of the golden banded skipper (Autochton cellus Boisduval and Le Conte, Hesperiidae) 
and an undescribed species of bee of the genus Dianthidium (Adanthidium) in the 
Megachilidae (Griswold, T. pers. com.). The skipper does not seem to be very specific and 
visits several species showing white flowers (Cortés-Palomec 2005), but the bee does ap- 
pear to only frequent Viola and it could be favoring gene flow between these two species. 
More specific studies of the pollinators would be needed to better understand their be- 
havior. 

In conclusion, we have shown that hybridization occurs between Viola grahamiiand 
V. hookeriana, and that the hybrids are morphologically distinct and intergrading from 
the parental taxa. Genetically, despite a larger similarity to V.grahamii, the hybrids have 
unique alleles. Ecologically, hybrids have a wider distribution and environmental toler- 
ance than any of the parental taxa, and at least in the area of study they are much more 
abundant than V. hookeriana. Similar flowering phenologies and effective pollen move- 
ment between the species has likely favored hybridization over a long period of time 
between the two species. 


1132 BRIT.ORG/SIDA 22(2) 


ACKNOWLEDGMENTS 


ACP would like to thank the Consejo Nacional de Ciencia y Tecnologia (CONACyTF 
Mexico) for financial support awarded under grant #128098. The authors also thank the 
Secretaria de Recursos Naturales y Pesca (Semarnap) for granting the collecting permits 
for our work. Thanks are also extended to Ricardo Wong and Juan Carlos Gonzales for 
providing field assistance and to Ross McCauley for help with data analysis. We would 
additionally like to thank Susan E. Yost, Nir L. Gil-ad, Mary J. Haywood, and Rebecca A. 
Peters for their useful comments and suggestions on this manuscript. 


REFERENCES 


Anoerson, E. 1954. An analysis of introgression in a population of stemless white violets. Ann. Mis- 
souri Bot. Gard. 41:263-269. 

Arcuibacp, JK, A.D. Wotre, and S.D. JoHNson. 2004. Hybridization and gene flow between a day- and 
night-flowering species of Zaluzianskya (Scrophulariaceae S.S., tribe Manuleeae). Amer. J. Bot. 
91:1333-1344. 

Arnoib, M.L. 1992. Natural hybridization as an evolutionary process. Annu. Rev. Ecol Syst. 23:237-261. 

Baitard, H.E., Jr. 1994. Violaceae. In: J. Rzedowski and G. Calderén de Rzedowski, eds. Flora del bajio y 
de regiones adyacentes. Fasciculo 31. Instituto de Ecologia, Centro Regional del Bajio, Patzcuaro. 

Battard, H.E., Jr.and D.E. Wuiex. 1994. Evidence for the recognition of Viola appalachiensis. Syst. Bot. 
19:523-538. 

Battarb, H.E, Jr, KJ. Syisma, and R.R. Kowal. 1999. Shrinking the violets: Phylogenetic relationships of 
infrageneric groups in Viola (Violaceae) based on Internal Transcribed Spacer DNA sequences. 
Syst. Bot. 23:439-458. 

Battaro, H.E., Jr. and KJ. Syrsma. 2000. Evolution and biogeography of the woody Hawaiian violets 
(Viola Violaceae): Arctic origins, herbaceous ancestry and bird dispersal. Evolution 54:1521-1532. 

Brainerd, E. 1924. Some natural violet hybrids of North America. Vermont Agric. Exp. Sta. Bull, 239:1- 
Zo: 

CALDERON DE R2FDOWSKI, G.2001.Violaceae.In:G. Calderon de Rzedowski, J. Rzedowski, and colaborators, 
eds. Flora Fanerogamica del Valle de México 2nd. ed. Instituto de Ecologia, A.C. and Comision 
Nacional para el Conocimiento y Uso de la Biodiversidad, Patzcuaro. Pp. 417-423. 

CLausen, J. 1964. Cytotaxonomy and distributional ecology of western North American violets. 
Madrono 17:173-204. 

Cortes-Patomec, A.C. 2005. Ecological factors, mixed breeding system and population genetic struc- 
ture in a subtropical and a temperate violet species. Ph.D. Dissertation. Ohio University, Athens. 

EvistRAND, N.C,,R. Writkus, and L.H. Riesserc. 1996. Distribution of spontaneous plant hybrids. Proc. Natl. 
Acad. Sci. 93:5090-5093. 

Excorrier, L. 1993. WINAMOVA. Analysis of molecular variance (Ver. 1.55).Genetics and Biometry Labo- 
ratory, University of Geneva. 

Gi-ap, N.L. 1998. The micromorphologies of seed coats and petal trichomes of the taxa of Viola 
subsect. Boreali-Americanae (Violaceae) and their utility in discerning orthospecies from hybrids. 
Brittonia 50:91-121. 

Haotey, E.B. and D.A. Levin. 1967. Habitat differences of three Liatris species and their hybrid deriva- 
tives in an interbreeding population. Amer. J. Bot.54:550-559. 

Harmaia, H. 2003.A deviating cytotype of Viola riviniana from Finland. Ann. Bot. Fennici 40:395-400. 

Hintze, J. 1999.NCSS 6.0 Statistical System for Windows. Kaysville 

James, J.K.and R.J. Assorr. 2005. Recent, allopatric, homoploid hybrid speciation: The origin of Senecio 


— 


CORTES-PALOMEC AND BALLARD ts 


squalidus (Asteraceae) in the British Isles from a hybrid zone on Mount Etna, Sicily. Evolution 
59:2533-2547. 
Jonsett, B., |. Nordat, and F.J. Roserts. 2000. Viola rupestris and its hybrids in Britain. Watsonia 23:269- 


Jupp, W.S.,C.S. Campett, E.A. KeLLoae, and PF. Stevens. 1999, Plant systematics: A phylogenetic approach. 
Sinauer Associates, Inc., Sunderland 

KRAHULKOVA, A., F. KRaHULeC, and J. KirscHNer. 1996. Introgressive hybridization between a native and an 
introduced species: Viola lutea subsp. sudetica versus V. tricolor. Folia Geobot. Phytotax. 31:219- 
244. 

Levin, D.A., J. FRANcisco-OrteGa, and R.K. JANsen. 1996. Hybridization and the extinction of rare plant 
species. Conservation Biol. 10:10-16. 

Manty, B.F.J. 1994. Multivariate statistical methods. Chapman and Hall, London. 

Marcussen, T.and L. Borcen. 2000. Allozymic variation and relationships within Viola subsection Viola 
(Violaceae). Plant Syst. Evol. 223:29-57. 

McCautey, R.A.and H.E. Bavaro, Jr. 2002. Inferring nativity and biogeographic affinities of central and 
marginal populations of Froelichia floridana (Amaranthaceae) from inter-simple sequence repeat 
(ISSR) markers. J. Torrey Bot. Soc. 129:311-325. 

Miter, M.P.1997.Tools for populati ic analysis (TFPGA) 1.3:A Windows program for the analysis 
of allozyme and molecular population genetic data. Computer Software available at 
www.marksgeneticsoftware.net. 

Neurrer, B., H. Auce, H. Mescu, U. Amarett, and R. Branoi. 1999. Spread of violets in polluted pine forest: 
Morphological and molecular evidence for the ecological importance of interspecific hybridiza- 
tion. Molec. Ecol. 8:365-377. 

O'Hanton, PC.,R.Peakait,and D.T. Briese. 1999. Amplified fragment length polymophism (AFLP) reveals 
introgression in weedy Onopordum thistles: hybridization and invasion. Molec. Ecol. 8:1239-1 246. 

Rryme, J.M.and D. Simecrtorr. 1996. Extinction by hybridization and introgression. Ann. Rev. Ecol. Syst. 
27:83-109. 

Rieseser, L.H. 1995. The role of hybridization in evolution: old wine in new skins. Amer. J. Bot. 82:944— 


RieseserG, LH. and N.C. Evistrano. 1993.What can molecular and morphological markers tell us about 
plant hybridization? C.R.P. Sci. 12:213-241. 

Russet, N.H. 1954. Three field studies of hybridization in the stemless white violets. Amer. J.Bot.41:679- 
686. 

Russet, N.H. 1955.Local introgression between Viola cucullata Ait.and V. septentrionalis Greene. Evo- 
lution 9:436-440. 

Vora,R.S.1988.A comparison of the spherical densiometer and ocular methods of estimating canopy 
cover. Great Basin Nat. 48:224-227. 

Wo re, A.D., Q.Y. Xianc, and S.R. KepHart. 1998a. Diploid hybrid speciation in Penstemon 
(Scrophulariaceae). Proc. Natl. Acad. Sci. U.S.A. 94:5112-5115. 

Wotre, A.D., Q.Y. XiAnG, and S.R. KepHart. 1998b. Assessing hybridization in natural populations of Pen- 
stemon (Scrophulariaceae) using hypervariable inter-simple sequence repeat (ISSR) bands. Molec. 
Ecol. 7:1107-1125. 

Wo re, A.D.and C.P.Ranoit.2001.Relationships within and among species of the holoparasitic genus 
Hyobanche (Orobanchaceae) inferred from ISSR banding patterns and nucleotide sequences. 
Syst. Bot. 26:1 20-130. 

ZAR, J.H. 1996. Biostatistical analysis. Prentice Hall, Englewood Cliffs. 


1134 BRIT.ORG/SIDA 22(2) 


Book REVIEWS 


Davip Fross and Dirter WILKEN. 2006. Ceanothus. (ISBN 0-88192-762-7, 978-0-88192-762- 
7, hbk.). Timber Press Inc, 133 $.W. Second Avenue, Suite 450, Portland, OR 97204- 
3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 
1-800-327-5680, 503-227-3070 fax.). $39.95, 272 pp., color photos, b/w illustrations, 
Teo 


As noted on the jacket: “Ceanothus, with so many plants that tolerate sun and shade, that thrive in arid conditions 


ae 
and thus conserve water, that bear a profusion of beautiful, fragrant flowers, deserves a book that will bring more 
of them into gardens and encourage greater appreciation of them in the wild, and this is that book.” Plenty of 


color photos of these spectacular shrubs and small trees. Statements of the geographic ranges are given; the only 
thing Lcan think of in improvement of this book, “The Compleat Ceanothus,” would] ave been generalized range 


Oo, 


maps. The text is divided into two substantial and readable sections (chapters in parenthesis): 


Ceanothus in the Garden and Landscape 


(Natural Distribution; History of Culti g Ceanothus, Disease and Pests; Cultivar Development; 
ee oes Ceanothus Species and C itats for an Garden and Landscape [94 pages| and Ceanothus 
Selec suide 

eer ns in th wis (an evoOnety and taxonomic overview) 


] = 
Distribution; E Orm anda Function in Ceanothus: ¢ 


les and descr intend and Other Ceanothus names) 


—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. 


David D. Stuart. 2006. Buddlejas. (ISBN 0-88192-688-4, 978-0-88192-688-0, hbk.). Royal 
Horticultural Society Plant Collector Guide. Royal Horticultural Society and Tim- 
ber Press Inc, 133 5.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. 
(Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327- 
5680, 503-227-3070 fax.). $34.95, 192 pp., color ne 7" x 9" 


A guid I Buddlej | | gard rith plenty of detailed information. “Buddlejas 


5 


are ihe most aocgmiadieoend eeneuing of garden pk ne ies acceptance of challenging soil conditions anc 
climate extremes and their resistance to pests (including deer, hares, and rabbits) are legendary.” The genus in- 
cludes about 100 species — the author lists and discusses 58 species of them (noting that he is concerned only 
with the woody species, vs. the herbaceous perennials). A middle s section provides beautiful color photos of most 
of the 58, along with many of the distinct cultivars of B. davidii. The super-hardy B. davidii (butterfly bush) is 


commonly used by breeders because of its abundant and nectar-rich flowers. “The major down side of the species 
is its ability to colonize inhospitable wasteland sites; the tiny winged seeds, blown far and wide, are capable of 
germinating wherever they settle, be it old tarmac, aioe or a crevice high on a wall. During World War II it 
besa known as the bombsite plant in Britain, as it was always the first plant to establish itself after an attack.” 

The North American natives B. marrubiifoli 1B. ulahensis are included in the book but not B. racemosa, B. 


scordioides, and B. sessiliflora. 

Chapter 1. Species 

Chapter 2. Hybrids and Cultivars 
Chapter 3. Care and Cultivation 

Chapter 4. Propagation 

Chapter 5. Wildlife Attracted to Buddlejas 


—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. 


SIDA 22(2): 1134. 2006 


HERBIVORY OF FERAL GOATS ON ESPIRITU SANTO ISLAND, 
GULF OF CALIFORNIA, MEXICO 


José Luis Leon-de la Luz and Reymundo Dominguez-Cadena 


Programa de Pl ion Ambiental y Conservacion 
G t At au ; R; logi dol NI. tf, (CIBNOR). 
Mar Bermejo 195, Col. Playa Palo de Santa Rita, 
La Paz, B.C.S. 23090, MEXICO 


jlleon04@cibnor.mx 


ABSTRACT 


Island fragil d iall luced ies, particularly large herbivores. In 
Mexico, ee are some dramatic cases. We ep alnated the influence of feral pone as well as the natives black 
jackrabbit and desert woodrat on the vegetation of Espiritu Santo Island, a land-bridge island in the southern 
part of the Baja California Peninsula. We found that goats consumed 36 vascular plant species, three times more 
species than natives jackrabbits and almost seven times more than woodrats. Also, goats consume all plant or- 
gans except roots, but reiterate browsing gradually affect vegetative and reproductive plant fitness. Our results 
support plans for the eradication of goats to spontaneously and progressively restore of the natural vegetation 
structure and composition. Further, a monitoring plan is recommended. 


Key Worps: goat herbivory, Capra hirca eradication, goat eradication, Baja California Islands 


RESUMEN 


Las islas son consideradas ecosistemas muy fragiles. Estas son sensibles a la introduccion de especies exoticas, 
icul herbi Fn Méxi ict | 44 EF te-trahoi laavebat, | 


if J 
la vegetacion en complejo insular Espiritu Santo, Baja California Sur, México, de las cabras silvestres contra el 
causado por las oe nativas, ra liebre negra y la rata de campo. Las cabras consumen 36 especies de plantas 
yeeculare es, tre mas esp que las HEDIES y casi 7 veces mas que las ratas. También, se encontr6 que las 
todo tipo d , pero el ramoneo reiterado afecta tanto la aptitud 


vegetativa y neprodilcnye de es pesca el -OS Eitedes apoyan el plan de erradicacion de las cabras con la 
finalidad de que espontanea y progresivamente se restaure la vegetacion en estructura y composici6n. 
Adicionalmente se requiere de un plan de monitoreo. 


INTRODUCTION 


Extinction and extirpation of native plants and animals from islands are significantly 
correlated with the introduction of exotic grazing mammals. Prevention of introduction 
and establishment of exotics on islands is vital, so that a program of eradication or con- 
trol of existing exotics is required on many islands around the world, and eradication of 
new introductions, as soon after Secccctne as possible, should be a high priority for con- 
servation natura by means of early aa and rapid 
response (McChesney & Tershy 1988; Burbidge 1999; enlan et al. 200 

Feral domestic goats (Capra hircus L.) are implicated in habitat ie and al- 
teration of species composition in sensitive insular ecosystems around the world 
(Scowcroft & Hobdy 1987; Jennings 1987; Parkes 1990). In the absence of population con- 
trols, goats become the ecologically dominant species on many islands, with the result 
that numerous native and endemic plant species have been extirpated or are threatened 
with extinction (Coblentz 1978; Keegan et al. 1994). A dramatic case is Isla Guadalupe in 
the Pacific Ocean near the state of Baja California, Mexico. This island was devastated y 


SIDA 22(2): 1135- 1143. 2006 


1136 BRIT.ORG/SIDA 22(2) 


grazing goats and erosion caused by soil trampling, to the extent that some native species 
and entire plant communities have vanished (Leon de la Luz et al. 2003; Oberbauer 2006). 

Espiritu Santo is a small archipelago of two major land bridge islands and several 
islets in the southwestern Gulf of California (Fig. 1). They are included in the Gulf of 
California island protection program covered by Mexican environmental laws, and ad- 
ministered by the Natural Protected Areas Commission (CONANP). The largest island is 
Isla Espiritu Santo is separated on the north by a 10 m wide channel from the second 
island, Isla Partida Sur. Additionally, there are the islets of La Ballena, El Gallo, La Gallina, 
and Los Islotes (Fig. 1). 

An undetermined number of domestic goats were introduced to Isla Espiritu Santo 
around the middle of the 20th Century to serve asa supply of fresh meat for local fisher- 
men. Currently, there are hundreds of feral goats on the island that are gradually destroy- 
ing the vegetation, eventually leading to extinction of native species. An informal ani- 
mal census by the authors in spring 2001 provided an estimate of 1,000 to 1,200 goats 
occupying Isla Espiritu Santo. Field observers could not be more precise because the gen- 
eral relief, escarpments, and steep slopes made conventional counting methods impos- 
sible. 

Because these islands are narrow and rocky, runoff is very rapid after the relatively 
rare rains. With little percolation into the soil, water is collects in occasional drainage 
pools. Thus, plants on the islands seem to have more profound water stress than on the 
main peninsula, a condition that is reflected by poor reproductive response, including 
less blooming, setting of fruit, and seed production than populations of the same species 
on the peninsula (author, pers. obs.). These conditions critically reduce the recovery ca- 
pacity of the plant community, and consequently, the dependent animal guilds. 

One of the greatest challenges for administrators of protected areas is maintaining 

biodiversity of native species and habitat loss. In Mexico, most of the islands have been 
protected to keep their biodiversity. This island, which is federal property and a protected 
area, is being considered for a program of goat eradication by the administrators 
(SEMARNAT, Baja California Sur). The administrators are familiar with the catastrophic 
effects of grazing activity on islands around the world and in Mexico. 
Based on field transects, this study compared and evaluated differential consumption of 
plants by feral goats and the endemic black jackrabbit (Le pus insularis Bryant) and desert 
woodrat (Neotoma lepida Thomas subsp. vicina Goldman) on Isla Espiritu Santo, com- 
paring their impacts on the vegetation. Following goat eradication, our data could pro- 
vide support for prediction of the recovery of the plant community. 


METHODS 


Site description 
The Espiritu Santo archipelago (24°24' to 24°36'N, 110°18' to 110°27' W) is part of the state 
of Baja California Sur. The nearest approach to the coast of the Baja California Peninsula 
is barely 8 km, northwest of the port at Pichilingue. The longest axis of the archipelago 
(NW-SE) is 19 km; the widest reach (E-W) is about 3.5 km. The archipelago covers 110 
km2, 87 km2 on Isla Espiritu Santo and 20 km? on Isla Partida. The highest point, near 
the midpoint of the big island is 540 m (Fig. 1). 

Geologically, the archipelago is composed of sedimentary rocks of volcanic origins 
(sandstones and conglomerates) of the Comondu Formation. Variations in resistance to 
weathering lead to a rugged topography. Except in a few areas, most of the surface has 


FERAL 1137 


110°20°W 


Los Islotes = 
x a’ GULF OF 
/ CH) CALIFORNIA 


Y 


Partida 


aa La Ballena 
/ poe, oe 7 


El Gallo a, aes 
La Gallina 2 oe ° 


ey a Espiritu Santo 


Pacific 

Ocean 2Kkm 
Fic. 1.6 g phi I] i f th + ly 1 tl ri Many: bth Calf o£ FA10L, Sa zs, A y 
over 200 m. C (+) indi he hia + at then tcl 1 (540 \ 


very steep rocky slopes where some soil is present beneath rocks. The uplands are table- 
lands where goats concentrate at nights. The west side of the island is highly dissected 
by ravines descending from the uplands to sea level. On the shore, salt flats form behind 
mangrove stands and sandy pocket beaches at the end of coves. The east side of the is- 
land is largely wave-cut cliffs and pebble beaches. 

Weather data are not available for the island, but an approximation, based on records 
from the city of La Paz, suggests annual mean rainfall <170 mm and potential evapora- 
tion of 2500 mm. Rainfall is erratic, with fall-winter-spring of 8 to 10 months without 
any rainfall. Occasional hurricanes in the late summer could drop up to three times the 
annual mean. 

Vegetation is typical of the peninsular Sonoran Desert, the sarcocaulescent scrubland 
(Shreve @ Wiggins 1964) is characterized by perennials with semi-succulent stems, in- 
cluding cacti and deciduous woody shrubs. According to the last inventory, 254 taxa are 
found (Rebman et al. 2002). Families include Compositae (28 taxa), Leguminosae (25), 
Euphorbiaceae (24), Graminae (20), and ee (15), these five cabae 44% of all the 
taxa. Some of the tarborescent forms f ically dominating the slopes 
are: “palo blanco” Lysiloma candida Brandegee, “salate” Ficus petiolaris HBK subsp. palmeri 
S. Wats., “cacachila” Karwinskia humboldtiana (Roem. & Sch.) Zucc., and “cardén” 
Pachycereus pringlei(S. Wats.) Britt. &@ Rose. 

Constant and strong winds noticeably contribute to lignosity in branches and a more 


1138 BRIT.ORG/SIDA 22(2) 


dwarflike appearance than examples of the same species growing on the Baja California 
Peninsula. With a north-south mountain ridge axis, the canyons run east and west, and 
the isolation generates different exposure-based plant associations on north-facing 
(shadow-side) and south-facing (sun-side) slopes. North-facing slopes having significantly 
more canopy and plant eon than the intensely exposed south-facing slopes. Sur Wor 
by the author, based on 400 m¢ * sampling plots, provide plant canopies oe 220m? 
on south-facing slopes and 305 m? on north-facing slopes, on opposite sides of the same 
ravine (author, unpublished data). 


ling 

In March 2001, ten straight-line transects of approximately 3 km each, were designed so 
that each would traverse several environments. Transects included beaches and man- 
grove (one site), arroyos and washes (three sites), slopes (four sites), and tablelands (two 
sites). Information recorded included: 1) plant and Olean consumed or injured, 2) evi- 
dence for the responsible herbi s species, based on characteristics of common sense, 
including type of bite on the plant and supporting evidence of droppings (pellets) beside 
the damaged plants. Droppings of goats are barrel form, approx. | cm long; those of jack- 
rabbits are elliptic-lenticular form, approx. 1 cm long, and those of desert woodrats are 
linear, approx. 1 cm long. 

Because the three herbivores typically defecate next to the browsed plant during feed- 
ing, we considered such occurrences related and definitive. Figure 2 shows a foraged plant 
and pellets of goats and black jackrabbits. Intensity of feeding activity was determined by 
the number of old scars on plants and the abundance and age of pellets around these plants. 


— 


RESULTS 
Table | assembles the information by sampled sectors, showing elevation, browsed spe- 
cies, consumed or injured plant organs, evidence of the herbivore species, and intensity 
of consumption based on scat abundance. From the transect surveys, we found that brows- 
ing by goats was concentrated on the more elevated mountains of the island near La 
Ballena and Los Candeleros Coves, and on uplands above +00 m 


Discussion 

In general terms, abundance and distribution of plants on the island is definitely depen- 
dent on terrain conditions, such as slope angle and exposure, as well as soil quality (soil 
deep and rockiness). These factors contribute to vegetation heterogeneity on the land- 
scape. We believe that many native plant species have been modified by goat browsing 
over the past several decades, knowing that goat herbivory directly affects fitness and 
reproductive success of mature plants and also contributes to consumption of juvenile 
plants. Donlan et al. (2002) documented positively changes in plant coverage on several 
Gulf of California islands after extirpation of exotic herbivores. 

Some actively browsed plant species are the perennial herbs Bebbia juncea (Benth.) 
Greene, Justicia californica (Benth.) Gibson, and Ditaxis lanceolata (Benth.) Pax & K. 
Hoffm. and the sub-shrubby Aeschynomene nivea Brandegee. We found that browsing of 
terminal branches promotes sprouting from basal nodes, leading to a branching archi- 
tecture of unusually short basal trunks and numerous branches. Several individual “palo 
blanco” Lysiloma candida, one of the few tree-like species, had trunks stripped of the high- 
tannin bark, as well as shrubby species of ‘lomboi” Jat ropha cinerea (Ortega) Muell-Arg. 


1 


ICANTCIA 


Sane, ©, 


foes 


Mt 


fst 


te 


oo 


hac) 


i 


la Pe 


1140 BRIT.ORG/SIDA 22(2) 


Taste 1. Perennial plant species affected by herbivory of feral goats, black jack-rabbits (BJR), and wood rats on 
Isla Espiritu Santo, Baja California Sur, Mexico, based in ten 3-km transects during a field survey in spring 2001. 


Plant species (asl) Consumed organ Herbivore Intensity 


Beaches and mangrove (0-2 m) 


Rhizophora mangle L. Leaves, Bark Goat Low 
Avicenia germinans (L.) L. Leaves, Bark Goat Low 
Monantochloe littoralis a Leaves JR Low 
Jouvea pilosa (Presl.) Scrib Leaves Goat Low 


Arroyos and washes (5-15 m) 


Viscainoa geniculata (Kell.) Greene Leaves Goat Low 
Forchammeria watsonii Rose Leaves Goat Low 
Euphorbia magdalenae (Benth.) Millsp. Buds Goat Low 
 aiadin candida Brandegee Flowers, Leaves, Bark Goat Low 
Melochia tomentosa L. Buds Goat Low 
sais ene (Engelm.) Gibson & Horak Stems BJR, Rat Low 
Is prin i (S.Wats.) Britt.& Rose Flower, Bark Goat, BJR Low 
rdsia chinensis (Link) Sch. Leaves oat Low 
Col ibaa viridis M. E. Jones Bark Goat Low 
Bebbia juncea (Benth.) G Leaves Goat ow 
ochemia poselgeri ener Britt.& Rose Stems BJR, Rat Hig 
Prosopis articulata S. Wats. Stems, Leaves Goat, BJR Medium 


Slopes (10-150 m) 


Justicia californica (Benth.) Gibson Stems, Leaves, Buds Goat High 
Ditaxis lanceolata (Benth.) Pax & K. Hoffm. Stems, Leaves Goat High 
Desmanthus fruticosus Rose Stems, Leaves Goat High 
ue nivea Brandegee Stems, Leaves Goat High 
Agave sobria Brandegee var. roseana (Trel.) Gentry — Flowers Goat High 
anes ia pacensis nas & carte Buds Goat High 
rtus (L.) Beauv.ex Roem. Leaves Goat Medium 
Cyiindropunts ia ciel a Weber Flowers, Fruit BJR, Rat Low 
Macrosiphonia hesperia |.M. Jhtn. Stems, Leaves Goat High 
Ficus petiolaris subsp. palmeri S. Wats. Leaves, Buds Goat High 
Bouteloua reflexa Swallen Leaves Goat Medium 
Solanum hindsianum Benth. Buds Goat Medium 


Tableland (150-450 m) 


cia Bases nee & Carte Stems Goat, BJR Medium 
bria Brandegee var. roseana (Trel.) Gentry Flowers, Leaves Goat Medium 
Buea eoinnata (Rose) Engler Buds Goat Medium 
Caesalpinia placida Brandegee Stems, Leaves Goat Medium 
Parkinsonia florida Benth. ex A. Gray Stems, Leaves, Buds Goat Medium 
subsp. peninsulae (Rose) Carter 
Fouquieria burragei Rose Buds Goat Medium 
latropha cinerea (Ortega) Muell.-Arg. Buds Goat Occasional 
hvaloua eandice Brandegee Bark Goat Medium 
Olneya tesota A. Gray Buds Goat Medium 
ea ia eis Engelm. Stems Goat, BJR Medium 
f hinensis (Link) Sch. Stem, Leaves Goat Medium 
Eushertia sneulenae (Benth.) Millsp. Buds Goat Occasional 
Supnorele DIOS (S.Wats.) Millsp. Buds Goat Occasional 


a (Benth.) G.L. Nesom Stems, Leaves Goat, BJR Occasional 


FERAL 1141 


Table 1. continued 


Plant species (asl) Consumed organ Herbivore Intensity 


Tableland (150-450 m) (cont.) 


Stenocereus gummosus (Engelm.) Gibson & Horak Stems BJR High 
Maytenus phyllanthoides Benth. Stems BJR, Rat Low 
Pedilanthus macrocarpus Benth. Stems BJR Occasional 
Opuntia cholla Weber Stems Goat, BJR Medium 
Hyptis laniflora Benth. Buds Goat, BJR Medium 
Pachycereus pringlei (S.Wats.) Britt. & Rose Stems Goat, BJR Occasional 
Echinocereus brandegeei (Coult.) K.Schum. Stems Rat Medium 


(see Fig 3). Some plants of “cardon” Pachycereus pringlei, a large columnar-arborescent 
cacti, have old scars on their trunks (>20-40 cm above the ground), evidence of brows- 
ing by goats and probably also jack-rabbits during periods of severe drought. Our obser- 
vations suggest that only two species of “choya” cacti (Cylindropuntia cholla Weber and 
G alcahes Weber) could benefit from goat browsing pecause sexual reproduction in these 


wo very limited and their jointed st asily, adhere to goat hair, and 
are easily dispersed and eventually they could settling by vegetative processes. 
Black jackrabbits and desert wood rats have a significantly lower impact on the veg- 


etation because, as smaller browsers; individually they consume less plant material and 
their population biomass seems to be less than the hundreds of goats. 

From the data assembled in Tables 1 and 2, we found that the most intense location 
of browsing by goats is on stems and, at least during spring, is frequent and intense. This 
browsing habit almost entirely consumes the floral and leaf shoots, the juicy and less 
woody tissues of the plants. After the rainy season, the goats concentrate on browsing 
the newly emerging foliage. 

The evidence indicates that a set of plant species display consumption of or injury to 
the meristematic apex of the branches, that is, the fresh and fluid-filled tissues, rather 
than woody tissue and stem bark. Clades of Opuntia tapona Engelm. are specially con- 
sumed by all the animals because the high water content. With freshwater in very short 
supply and stored in “tinajas” after rainfall, goats manage to prosper by drinking salty 
water (Burke 1990). During our field work, we observed a few goats drinking seawater at 
the beach at sunset, which suggests daily or close to daily migration from the uplands to 
the shore. At the shore, goats also consume halophytic plants, including mangrove and 
species of salt grasses. 

Our findings demonstrated that goats are the most significant herbivores, capable of 
consuming almost all plant parts. While the native black jackrabbit and desert woodrat 
have the same feeding habits as goats and inhabited the island for millennia, their den- 
sity and biomass have far less impact on the vegetation. 

Finally, there was no direct evidence that goats consume an entire plant; however, 
high intensity herbivory strongly affects the fitness of each grazed plant, and can even- 
tually lead its death. We conclude that the natural balance of vegetation on Isla Espiritu 
Santo is deleteriously affected by goat herbivory, which combined with the impact of the 
native foragers, increases the deleterious effects of browsing on a large group of plant 
species. Our data supports plans for the total eradication of goats from the archipelago. 


BRIT.ORG/SIDA 22(2) 


Fic. 3. An exemonplar of Jat, } I | I t te hark 1 brok I } 

r f J ~ ad re i 
Taste 2.Concentrated data of plants consumed by herbivores on Isla Espiritu Santo, Gulf of California, Mexico. 
Herbivore Plant species Plant organs Sector of activity 

consumed consumed 

Goats 36 Flowers, buds, leaves, young stems, bark Slopes, high tablelands 
Jackrabbits 13 Stems Arroyos, high tablelands 
Wooadrats 5 Stems, fruit Arroyos, slopes 


On an ecological basis, black jackrabbit and woodrats are foragers that control the veg- 
etation growth. 

After eradication a monitoring plan is strongly recommended. We hope that the same 
plants indicated in Table | will increase their density and size. In fact, eradication of goats 
is currently underway by the administrators of the island. The project is expected to be 
completed by winter 2006/07, with only radio-collared and sterile sentinel goats remain- 
ing (Donlan et al. 2002). 


ACKNOWLEDGMENTS 
We appreciate the English editing of Ira Fogel at Centro Investigaciones Bioldgicas del 
Noroeste, as well Miguel Dominguez for field expertise during our surveys of the island. 


FERAL AND 1143 


REFERENCES 


Bursince, A.A. 1999.Conservation values and management of Australian islands for non-volant mam- 
mal conservation. Australian Mammalogy 21:67-74. 

Burke, M.G. 1990. Seawater consumption and water economy of tropical feral goats. Biotropica 22: 
416-419. 

Costentz, B.E. 1978. The effects of feral goats (Capra hircus) on island ecosystems. Biol. Conservation 
13: 279-286. 

Dontan, C.J., B.R. TersHy, and D.A. Croit. 2002. Islands and introduced herbivores: conservation action 
as ecosystem experimentation: applied issues with predators and predation. J. Appl. Ecol. 39: 


JeNNincs, G.Y. 1987. Guadalupe: An island refuge under siege. Oceans 20(5):40-45. 

Keecan, D.R., B.E. Costentz, and C.S. WincHEeLL. 1994. Ecology of feral goats eradicated on San Clemente 
Island, California. In: Halvorson, W.L., GJ. Maender, eds. The Fourth California Islands Symposium: 
update on the status of resources. Santa Barbara Museum of Natural History, Santa Barbara, CA. 
Pp. 323-330. 

Leon ve ta Luz, J.L., J.P. Resman, and T. Osersauer. 2003. On the urgency of conservation on Guadalupe 
Island, Mexico: Is it a lost paradise? Biodivers. Conservation 12:1073-1082. 

McCuesney, G.J.and B.R. TersHy. 1988. History and status of introduced mammals and impacts to breed- 
ing seabirds on the California Channel and northwestern Baja California Islands.Colon.Waterbirds 
21:335-347. 

Osersauer, T.A.La vegetacion de isla Guadalupe. Entonces y Ahora. 2006. In: K. Santos del Prado and E. 
Peters. Isla Guadalupe: Restauracion y Conservacion. Instituto Nacional de Ecologia (INE- 
SEMARNAT), México. Pp. 39-53. 

Parkes, J.P. 1990. Eradication of feral goats on islands and habitat islands. J. Royal Soc. New Zealand 
20:297-304, 

Resman, J.P, JL. LEON bE LA Luz, and R.V. Moran. 2002. Vascular plants of the Gulf Islands (Appendix 4.1). 
In:T.J. Case, M.L. Cody and E. Ezcurra, eds. A new island biogeography of the Sea of Cortés. Oxford 
University Press. New York. Pp.465-512. 

Scowcrort, P.G.and R. Hospy. 1987.Recovery of goat-damaged vegetation in an insular tropical mon- 
tane forest. Biotropica 19:208-215. 

SHreve, F.and ILL. Wicains. 1964. Vegetation and flora of the Sonoran Desert. Stanford University Press, 
Stanford, CA. 


1144 BRIT.ORG/SIDA 22(2) 
Book REVIEWS 

Doucias D. Stokke and Leste H. Groom (ed.). 2006. Characterization of the Cellulosic Cell 
Wall. (ISBN 0-8138-0439-6, 978-0-8138-0439-2, hbk.). Black well Publishing. 2121 State 
Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK 
(Orders: orders@ames.black wellpublishing.com, 515-292-0140, 515-292-3348 fax I- 
800-862-0657, www.blackwellprofessional.com). $149.99, 274 pp., color photos, nu- 
merous b/w illustrations, 7" X 10' 


ing ... but so a more. This collection of 


(oy 


“Crack” as the twig breaks, the slightly elastic wood suddenly yielc 


studies, fundamental and applied science, represents the proceedings of a work : sored by the USDA 
Forest Service, the Society ‘ Wo a Science and Technology, aus lowa State U niversity — “the eeu i collabora- 


tion between three institutions witha ae: history of res g lignoc cel- 


lulosic substances, primarily wood.” The book is “intended for scientists, university faculty, sec trte students, 
and applied researchers in the fields wood science and technology, cellulose science, and biomaterials.” The 
book is arranged in three sections: 

Cell Wall Assembly and Function: New Frontiers (5 chapters, e.g. tracheid and sclereid differentiation in callus 
cultures of Pinus radiata; mechanics of the wood cell wall; prediction of wood structural patterns by using eco- 


logical models of plants water relations) 


Probing Cell Wall Structure: Advances in Analysis (6 chapters, e.g: determining xylem cell wall properties by us- 


ing model plant species; near infrared spectroscopic monitenng of the diffusion process of deuterium-labeled 


molecules in wood; wood stiffness by X-ray diffractometry) 
Mesostructure dnd Applications: Science in Practice (8 chapters, e.g: variation in kink and curl of longleaf pine 
fibers; wood structure and adhesive bond strength). 


—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


Marti J. GRoom, Gary K. Merve, and C. RoNALD CARROLL. 2005. Principles of Conservation 
Biology, Third Edition. (ISBN 0-87893-518-5, hbk.). Sinauer Associates, Inc., 23 Plumtree 
Road, Sunderland, MA 01375-0407; U.S.A. (Orders: orders@sinauer.com, 
wwwsinauer.com, 413-549-4300, 413-549-1118 fax). $92.95, 699 pp., 369 illustrations, 
S172 Sr 


This is a major update and revision of the 1997 Second Edition, with much new information and guided by a 


survey of “over 60 instructors on their needs for teaching conservation biology.” The text is divided into three 


major units: Conceptual Foundations for Conservation Biology (5 chapters); Focuson a yl hreatsto Biodiversity 
] t 


(6 chapters), Approaches to Solving Conservation Problems (7 chapters). A “companion sa variety 
of study materials and supplemental resources to accompany the textbook.” On the first page of Chi ices I: “the 
cumulative impacts of 6.4 billion people (Figure 1.1) have stressed the many ecological support systems of the 
planet. ... All prope should neCOB Ize the degree to irae human impacts affect the natural world, and in turn, 


| } al 


diminish our biologists conservation should have this book at hand (or 


one like it, if there’s anther: so seed) —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort 
Worth, TX 76102-4068, U.S.A. 


SIDA 22(2): 1144. 2006 


A DEMOGRAPHIC AND ECOLOGICAL ANALYSIS OF 
GEOCARPON MINIMUM (CARYOPHYLLACEAE): A 
FEDERALLY THREATENED SPECIES IN SOUTHWEST MISSOURI 


Tim Smith! Joseph S. Ely 


Missouri ee Department of Biology 
PO, Central Missouri State University 
Jefferson City, iaissonan see 0180, U.S.A. Warrensburg, Missouri 64093, U.S.A. 
ABSTRACT 
A Geocarpon minimum population growing on a sandstone glade in Dade pei! Missouri was monitored for ten 
years (1994-2003). Plants were counted annually in eight permanent plots at two scales (0.01 and 1.0 m2) to determine 


population fluctuations through time. Soil depths as well as cover of eee ce lichens, vascular plants, and bare 
soot were recorded in subplots between 1994 and 2000 to determine any ae association with G. minimum 
mbers. The number of G. minimum plants in the eight plots varied from 1 to 4230, a mean of 1332. Mean soil 


— 


aa of subplots containing G. minimum was 19 mm. There was a weal association (1, = “0 16, P<0O. ool) seubibidae soi 
depth and Geocarpon density. Regressic e 
soil depth to about 20 mm and ee ee In addition, there were significant differences among vegetation 


- al 


that G 


types (P < 0.05) with the exception of lichens (P > ac There was a ag pious difference between G. minimum 
W 


population numbers and year (P < 0.05). There was 1 scale and the Spearman rank correlations, 


a 
however, when ecological Broups were cgunpined into oie eatteoas (ie. lichens and bryophytes were lumpe 


into 5), patterns E bar are tock areas indicates a dynamic process 
] Be | a ]; 


of soil oT eae into Banc out of plots, | 


lar pla nte s Hee | gay I ] y ee i ] a a | f= 


crease In cove ies of moss, Vasc u- 


disturbance and with minimal competition from pb onaytes or perenaal vasa plants Although seed kau was 


L I 
{.]] ] 4 
LOLLOVY ing yeAalo W ith VII tually 


] 


node Teeter hi : 
of 


RESUMEN 


Una Poe de soles ble i m is mopireraca cuante ae anos caeaiipicns enun nenoueente bas 


aL enisca 
y 
numero de pate en ar Reach durante el periodo de estudio, las plantas se contaron anualmente en ocho 
parcelas permanente las (0.01 y 1.0 m2). Durante los anos 1994 hasta el 2000 fueron evaluados 
datos sobre ee de stele coe de briotitas, liquenes, plantas vasculares y area de tierra o rocas sin 
q P y 


plantas en subparcelas con el { ito ded la posible asociacion de estas ae con el numero de 
plantas de G. minimum. El numero de ae de G. minimum en las ocho parcelas vario de 1 a 4230 con un 
promedio de 1332. La oes poe del suelo eeu las ven con G. minimum fue “ 19 mm. La 
asociacion entre la y densidad de pl ‘pon fue minima (r, = 0.16, P < 0.001). El 
andalisis de regresion demostr6 que el numero de G. minimum parece aumentar a Rene re que la profundidad de 
suelo alcanza los 20 mm y disminuye a mayor profundidad. Ademas, la diferencia entre tipos de vegetacion (P< 


0.05), con excepcion de los liquenes (P >= 0.05), fue significativa. La diferencia fue significativa también entre el 
numero ue oe de G. minimumy el ano del muestreo (P < 0.05). Se observ6 una discontinuidad entre escala y 
le rango de Spearman, sin embargo, al combinar los grup oldgi n categorias mayores 


(por ee los aie y las pbloritas se unieron en e eatseCHe de las criptgaas) los patrones fueron mas 
claros a través de las escalas. La as indica la existencia de un 
proceso dinamico de movimiento del suelo dentro y fuera de las parcelas, pero is ne sobre cobertura no indi- 


can un crecimiento constante eae Selo es eos pleats eo o liquenes durante los diez anos del 
} -] 


estudio. I - tienen 


Acie ell 
i & A Fig L oO 


'Tim Smith, corresponding author: Telephone: (573) 522-4115; Fax: (573) 526-5582; E-mail: tim.smith@mdc.mo.g 


SIDA 22(2): 1145- 1157. 2006 


1146 BRIT.ORG/SIDA 22(2) 


en este estudio, su importancia fue evidente. En base a los datos anuales, esta poblacion de G. minimum tiene la 


capacidad ae reproducirse considerablemente en anos siguientes aunque no exista produccion de semilla en 


anos anterio 


INTRODUCTION 


Geocarpon minimum Mack. (Caryophyllaceae) isa tiny, succulent winter annual that grows 


on sandstone glades in Missouri, on saline soil prairies in Arkansas and Louisiana (U.S. 
Fish and Wildlife Service 1993; McInnis et al. 1993; Orzell & Bridges 1987), and on saline 
barrens in Texas (Keith et al. 2004) (Fig. 1). The diminutive height of the plant (up to 4 cm), 
its inconspicuous flowers, and its limited distribution are probably responsible for it not 
having been described until 1914 (Mackenzie 1914). Taller plants are usually multi-stemmed 
and many-flowered. The smallest plants are only a few mm in height and produce just one 
flower. Both morphologies are often present in a single population, requiring surveyors to 
acquire a broader search image to locate the full range of plant forms. Plants turn froma 
whitish-green to wine-colored to light brown as they mature and senesce. 

Geocarpon minimum was designated by the U.S. Fish and Wildlife Service as a feder- 
ally threatened species in 1987 because of low population numbers (U.S. Fish and Wildlife 
Service 1987). It is known today from fewer than 35 sites (Nienaber 2005). Twenty-four ex- 
tant Missouri populations are known from Pennsylvanian-age sandstone glades in seven 
southwestern counties, giving it a state rank of S2 (Missouri Dept. of Conservation 2006). 
The sandstone glades occur in 1 the Springfield Plain subsection of the Ozark Highlands 
ecological section and ar idered imperiled (a rank of S2) because of their rarity within 
the state (Nelson 2005). G. minimum is considered extirpated from its type locality, Jasper 
County, Missouri, where suitable habitat may have been lost due to land-use changes. 

Prior to this study, little was known about the management requirements for main- 
taining populations of G. minimum or the range of year-to-year population fluctuations. 
Several sites for the species occur on public lands where lack of information has led toa 
lack of management action. There was concern that some periodic disturbance might be 
necessary to maintain suitable microhabitat for G. minimum. On many of the glades where 
the species occurs, the thin soil near the rock/soil interface is dominated by dense mats 
of moss, except where trampling, off-road vehicle use, or water flow have maintained 
areas of loose sand. The purpose of this study was to gather long-term monitoring data 
that could provide needed insights for effective management of G. minimum populations. 
Specific objectives of this study were to: 

1) provide a long-term census of G. minimumat the Flint Hills Glade site, 

2) determine the correlation of bryophytes, lichens, vascular plants, and soil depth 
on the G. minimum population, 

3) provide management recommendations for the conservation of this species. 


MATERIALS AND METHODS 


Study Site 

The study site, Flint Hill Glades, is located in Dade County, Missouri, near Stockton Lake, 
on land owned and managed by the U.S. Army Corps of Engineers (Fig. 2). Geocarpon 
minimum was discovered at Flint Hill Glades during a 1989 survey for the species 
(Thurman & Hickey 1989). The site is atypical in one respect; the impoundment of Stock- 
ton Lake inundated the creek valley below the glade and floods the lower portions of the 
glade during extremely high lake levels, which occurred during the summer of 1993. 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1147 


Fic. 1. Geocarpon minimum plant. Photo by Rick Thom 


It is estimated that less than five-percent of the six-acre glade area is suitable habitat 
for G. minimum, which grows in greatest abundance in shallow sand- and pebble-filled 
depressions in the sandstone bedrock that are often devoid of other plants (Fig. 3). Based 
on research at another Dade County site, Morgan (1986) reported that optimum habitat 
was found in zones of shallow soils (1-5 cm) over sandstone bedrock. These microhabi- 
tats are maintained by flow of water over the bedrock that deposits sand and pebbles 
into the depressions. Water flow can also carry away previous deposition and prevent 
lichen or vascular plant succession. An intermittent drainage flows through the study 
site carrying runoff from adjacent grazed pasture on private property into Stockton Lake 
at the lower edge of the glade. 

Much of the surface bedrock is covered by a thick mat of mosses (principally 
Polytrichum ohiense Ren. & Card.) intermixed with lichens, liverworts, and a few vascu- 
lar plant species. Areas of deeper soil within the glade support grasses, shrubs (e.g. 
Vaccinium arboreum Marshall) and dwarfed trees (e.g. Quercus marilandica Muenchh. 
and Amelanchier arborea (FE Michx.) Fernald). Geocarpon minimum plants were observed 
in the grazed portion of the glade on adjacent private land during the course of this study. 
They are found only near a bedrock exposure where pasture grasses are not established. 
The dense beds of moss found on the study site are not present in the pasture. The tram- 
pling from cattle may help to maintain the loose sand particles that accumulate to form 
G. minimum microhabitats. The pasture area may be a source of G. minimum seed for the 
monitoring area. No monitoring plots were tablished in the pasture, primarily because 
cattle grazing is not viewed asa realistic management tool for most publicly-owned sites. 


Data Collection 
Ten plots were established at the site in April 1994. The 1.0 m x 1.0 m quadrats were sub- 


1148 BRIT.ORG/SIDA 22(2) 


, Stockton Lake, Dade County, Missouri 


jectively located in areas containing G. minimum in close proximity to areas dominated 
by mosses, lichens, or perennial vascular plants. Two opposite corners of each plot were 
marked by masonry nails driven into holes that were drilled into the bedrock. 

Two of the original ten plots eventually were dropped from the study because they 


— 
— 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1149 


Fic. 3. Habitat at Flint Hill Glades. Photo by Tim Smith. 


did not support any G. minimum after 1994. Inundation of part of the glade in 1993 cre- 
ated temporary habitat in these plots due to silt and sand deposition with little vegeta- 
tive competition. At one plot, sand and silt was deposited into a patch of bryophytes on 
bedrock, temporarily providing habitat that supported G. minimum. The bryophytes had 
been killed by the prolonged inundation and were soon gone, leaving only bare rock at 
subsequent surveys. The other plot was located near the lake shore and included essen- 
tially bare sand after the inundation, although it had deep enough soils to support pe- 
rennial vascular plants. Although G. minimum was found in the plot when it was estab- 
lished in 1994, this plot thereafter became covered with debris from subsequent rises in 
the lake level and no longer contained exposed sand or G. minimum. 

Alm X 1m (1.0 m?) wooden frame was used to monitor the permanent plots. Fly- 
fishing line was strung on the inside of the quadrat to further subdivide the 1.0 m2 plots 
into 100 subplots of 10 cm x 10 cm (0.01 m2). The number of G. minimum plants was 
recorded in each subplot. Soil depth was measured at the center of each subplot by using 
a nail to probe to bedrock and then measuring the length of the nail that had entered the 
soil. Bare rock was recorded as zero soil depth. Vegetative cover was estimated using seven 
cover classes (a modified Daubenmire cover class) and dominant cover was character- 
ized as mosses, lichens, vascular plants or bare rock/soil (Daubenmire 1959) The seven 
cover classes were: 1 (O-1%), 2 1-5%), 3 (5-25%), 4 (25-50%), 5 (50-75%), 6 (75-95%) and 7 
(95-100%). 

In the spring of 1995, three additional plots were established within the study area, 
in moss-dominated areas that did not contain G. minimum. These plots were created by 
removing a 1m X 1m section of dense moss to expose underlying mineral soil or bed- 


1150 BRIT.ORG/SIDA 22(2) 


2500 


2000 


1500 


Numbers of plants 


1000 


1994 
1995 
1996 1997 


1998 4999 
ae 2000 
Monitoring year 2001 5n92 


2003 


Fic. 4. Numbers of Geocarpon minimum plants i lots from 1994 to 2003. 


rock. These three plots were subsequently monitored for G. minimum, but soil and cover 
measurements were not taken. 


Data Analysis 

The subplot (0.01 m2) values of G. minimum numbers, soil depth, and cover classes for 
each vegetation type (see above) were summed within each large plot (iec., 1.0 m*) and 
were subsequently used for all analyses (Spearman rank correlation, analysis of vari- 
ance, and regression analyses). The independent variables were year, soil depth (mm), 
and cover values of bryophytes, lichens, and vascular peu Lichens and bryophytes 
were bined into a single category (cryptogams) and vascular plants and cryptograms 
were also combined into a single category of total vegetation. 

Spearman rank correlation coefficients, rs, (Zar 1999; Sokal &@ Rohlf 1995) were cal- 
culated on both subplots and large plots in order to determine strength of association 
among G. minimum, year, soil depth, bryophytes, lichens, cryptogams, vascular plants, 
and total vegetation at two spatial scales (0.01 m2 and 1.0 m2). In addition, partial correla- 
tions were also determined on ranked data controlling for the effect of year to determine 
potential associations among the vegetation types and soil. 

Model Il regression analyses were performed with the number of subplots that con- 
tain G. minimumas the dependent variable and soil depth as the independent variable. 
The regression analyses were performed on subplots only. Note that both soil depth and 
the number of subplots that contain G. minimum follow a normal distribution, as indi- 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1151 


cated by a Kolmogorov-Smirnov normality test (P > 0.05). Specific regression analyses 
were performed independently on soil depth range of 1 to 20 mm and 21 to 110 mm. The 
justification for the latter was to show that the subplots containing G. minimum increase 
with increasing soil depth to 20 mm and declines thereafter. 

A Friedman repeated measures (FRM) analysis of variance (ANOVA) (a non-para- 
metric equivalent to the parametric repeated measures ANOVA) was used to determine 
differences among population numbers annually. Natural log transformed G. minimum 
numbers was the dependent variable, year was the repeated measures factor, and plot 
was the subject factor (Zar 1999; Sokal & Rohlf 1995). Because some of the plots had zeros 
and ones, it was necessary to add a count of 2 (a constant) to each observation for natural 
log (in) transformation (Zar 1999; Sokal & Rohlf 1995). The transformation was done in 
order to improve statistical models (Quinn & Keough 2002) and to maximally normal- 
ize the data. Independent variables, with the exception of year and plot, were treated as 
random in all models because they were not specifically controlled (Zar 1999; Sokal & 
Rohlf 1995). The year 1996 was excluded from all analyses because only one G. minimum 
plant was observed for that year and thus it was considered as an outlier. 

Additional FRM ANOVAs were performed on cover classes of bryophytes, lichens, 
vascular plants, cryptogams, total vegetation, and soil depth. There were three cover classes 
for bryophytes, lichens, vascular plants, cryptogams, and total vegetation while soil depth 
was placed into six. FRM ANOVA was performed on the large plots only. Independent 
variables were natural log transformed as needed. Additional repeated measure ANOVAs 
(Kruskal-Wallis) were performed without 1999 data in order to ascertain the relation- 
ship between soil depth and vegetation groups without the possibility of an interacting 
effect or confounding influence for the year 1999. The year 1999 was considered an out- 
lier similarly to 1996 but with large numbers of Geocarpon observed for that year. The 
alpha level for all analyses was 0.05. 


RESULTS AND DISCUSSION 


Population Fluctuations 
Winter annual glade plants commonly exhibit annual fluctuations in population size. 
Available moisture for fall germination and spring growth can vary greatly, with low 
moisture making shallow-soiled areas hostile for plant growth. Long-term monitoring is 
important in order to document the range of fluctuations in population size. Knowledge 
of this range improves our ability to interpret population data from sites where long- 
term data are not available. The ability of a species’ seed bank to repopulate a site after a 
poor seed production year has obvious implications for long-term conservation. 
Results of annual G. minimum counts in eight plots are depicted in Figure 4. The 
cumulative annual totals from all eight plots ranged from one plant (1996) to 4230 plants 
(2003). Although all of the plots are the same size, the amount and quality of the G. mini- 
mum habitat within each plot varies, with some plots containing more bare rock or denser 
vegetative cover. Plots 6 and 7, which often contained the most Geocarpon, provide a 
greater percentage of thin soil with little competition from bryophytes, lichens, or vas- 
cular plants. The small number of G. minimum plants at the site in 1996 was probably 
due toa prolonged dry period during early spring, prior to monitoring conducted in April. 
Only one additional G. minimum plant was observed outside of the plots during that 
year’s monitoring. More substantial numbers of G. minimum were found at other sites in 
the region that spring, indicating the critical effect of very local drought on the popula- 


1152 BRIT.ORG/SIDA 22(2) 


tion at the study site. No insect herbivory of G. minimum was observed at the study site 
or at any other Missouri sites. There was a moderate positive correlation (1; = 0.44, P< 
0.001) of G. minimum with year at the 1.0 m? scale (Table 1). Plot to plot variation was 
significant between 1997 and 1999 (RM: X? = 1L7, P = 0.045; Tukey: q = 4.16, P < 0.05). 
Although the population fluctuated from year to year depending on local conditions, 
there was an overall increase in the population over the years. 


Soil Depth Considerations 

The stream and sheet flow of water over the glade due to precipitation creates a dynamic 
environment for G. minimum, with the depth and location of the loose, sandy soil with 
small pebbles changing dramatically through time. It was not uncommon to find G. mini- 
mum plants that had been uprooted and probably displaced by water and soil movement. 
It seems likely that the succulent stems are at least somewhat tolerant of displacement 
and can become established at new locations, especially when partially buried in sand. 

The mean soil depth for subplots containing G. minimum was 19 mm (Fig. 5a, b). In 
comparison, Somers et al. (1986) found the shallowest soil depths on xeric communities 
of the limestone glades of Middle Tennessee to be (35 + 17 mm), with such areas domi- 
nated by annuals and mosses, including several endemic species. In this study, soil depth 
was measured at the center of each subplot rather than at the actual location of any G. 
minimum plant. Therefore the high number of subplots with zero soil depth does not 
imply that plants were growing on bare rock but rather that they grew within about 7 cm 
(the distance from the center to a corner of a subplot) of the bare rock/soil interface. 
Nevertheless, there was a significant association between soil depth and G. minimum 
numbers (rs = 0.16, P< 0.001 at 0.10 m? scale; r;= 0.41, P < 0.001 at 1.0 m2 scale)(Fig. 5b 
and Table 1). Note that the Spearman rank correlation between soil depth and the fre- 
quency of subplots containing G. minimum was -0.46 (Fig. 5a). It was observed that in 
areas of deeper soil, perennial vascular plants were more often present. Where vascular 
plants become numerous, fallen leaves from trees can lodge between stems and accumu- 
late, whereas G. minimum-rich areas of the glade are windblown and/or washed by mov- 
ing water and remain largely free of leaves. 

Shallow soil, per se, is not a requirement for G. minimum growth. Rather, it is the 
mechanism by which plant competition is minimized on the sandstone glades. This was 
borne out by the growth of G. minimum on deeper soils following removal of other veg- 
etation and litter by prolonged inundation and wave action on the lower portions of the 
study site in 1993. This is also apparent with the observation that G. minimum grows on 
deeper soils in the saline prairies of Arkansas, Louisiana and Texas, where high sodium 
levels prevent the growth of competing vegetation. In addition, there were significant 
differences with soil depth and the distribution of bryophytes, lichens, and vascular plants 
at the 0.01 m? scale as indicated by the Kruskal-Wallis analysis of variance and/or the 
Spearman rank correlation coefficients (bryophytes: H4 = 19.96, P < 0.001, rs = -0.12, P< 
0.001; lichens: H4 = 19.10, P = 0.002, r; = -0.110, P > 0.05; and vascular plants: H4 = 17.18, P= 
0.002, rs = 0.445, P < 0.001)(Fig. 5c, d, and Table 1). 

Both soil depth and the number of subplots that contain G. minimum passed the 
normality test (kolmogorov-Smirnov P > 0.05). The results of the model Il regression 
analysis show that G. minimum increases with increasing soil depth until about 20 mm 
(F2, 18 = 4.45; P < 0.05) albeit with a low adjusted r? (15.4%). The regression model ac- 
counted for fifteen percent of the variation and is consistent with the results of the 
Spearman rank correlations (Fig. 5a, b and Table 1). An additional regression analysis 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEQCARPON MINIMUM IN MISSOURI 1153 
Taste 1.Spearman rank correlation coefficient for Yea ini bryophytes, lichens, vascular plants, 
cryptogams, total vegetation, and soil at the ane 0. 01m: 2, n = 862, (above the diagonal) and large plot (1.0 
m’),n = 50 (below diagonal) scales. Correlations between large plot and subplot data (along the diagonal) 
were not calculated because the data are not paired. * = significance at the 0.05 level. 


Year G.minimum  Bryophytes _Lichens Vascular Cryptogam _sTotal Soil 
Plants Vegetation 

Year _ 0.440* “0.23” -0.317* 0.022 0331" SO: 53i4" 0.038 
G. minimum 0.190 = -0.230* -0.168*  0.186* -0.305* -0.236* 0.162* 
Bryophytes -0.161 -0.098 - -0.160*  -0.577* 0.729% 0.362* -0.118* 

i -0.184 -0.045 0.367* - -0.442*  0.550* 0.298* -0.027 
Vascular Plants -0.024 0.204 -0.154 -0.273 ~ -0.781* 0.070* 0.165* 
Cryptogam -0.283*  -0.099 0:727* 0.829% -0.346* - 0.542* -0.111* 
Total Vegetation -0.312*  -0.041 0.663* 0.755* 0.120 0.871* - 0.043 
Soil 0.042 0.408* -0.199 -0.256 0.471* = -0.222 0.448* - 


shows that G. minimum numbers decrease with increasing soil depth beyond 20 mm (F2, 
gg = 68.45; P < 0.0001) with an adjusted r? of 43.1% (Fig. 5a, b). These results suggest that 
G. minimum numbers increase with increasing soil depth where CoMpe Non isata mini- 
mum but decreases thereafter. The latter observation is probably due to th petition 
from perennial vascular plants, which are more prevalent on deeper soils. 


Vegetation Correlations 

Large areas of the glade’s sandstone bedrock are covered with dense bryophytes and it 
was expected that cover measurements would demonstrate that bryophytes were spread- 
ing into G. minimum populations. Although bryophytes were negatively correlated with 
G. minimum numbersat the subplot scale (1; = -0.23, P < 0.001), they were not at the large 
plot scale. Similar results were observed with lichens and G. minimum numbers in that 
there was a negative relationship at the subplot scale but not at the large plot scale (Table 
1). Although there is some discontinuity among scale with the latter two groups, there is 
a strong negative association (r; = -0.78, P < 0.001) between cryptogams (combined cat- 
egory of bryophytes and lichens) and G. minimum numbers (Fig. 5f and Table 1). The 
latter may suggest that cryptogam competition is a factor in depressing G. minimum 
populations. 

A positive association occurred with coverage by vascular plants and G. minimum 
(r5= 0.19, P< 0.001) at the small plot scale but not at the next scale (Fig. 5e and Table 1). It 
may be that through time G. minimum has persisted only on those areas of the glade that 
are subjected to the same periodic disturbances that make it difficult for mosses and vas- 
cular plants to become established. Lichen cover declined steadily from 35.1% in 1994 to 
10.9% in 1998. Perhaps this aspect of the monitoring would show better statistical trends 
if conducted over a longer period and with more detailed data collection with respect to 
the species of vascular plants and cryptogams across population sites. Certainly the 
movement of soil over the bedrock would explain the erratic variation in numbers of 
subplots dominated by bare soil or bedrock. There are similarities between G. minimum 
population dynamics and winter desert annuals in that there are large population num- 
ber changes from year to year. Guo et al. (2002) discussed dramatic fluctuations in abun- 
dance, productivity, and diversity among desert winter annuals. Furthermore, the popu- 
lation dynamics of Vulpia ciliata Link. has been well studied (Watkinson et al. 2000; 
Watkinson 1990; Watkinson & Davy 1985; Watkinson & Harper 1978). This annual grass, 


1154 BRIT.ORG/SIDA 22(2) 


¢ ] 60 | . < 1 
2 & ie a | € | - 1, = 0.16, p< 0.00 b- 
2 & | & 50 | | 
Qa iS) 3 il e° 
28 40 E 40 
a © 5 
5 oD 30 ° | 
2 . | = 0 | 
c 
8 g 20 & 20 
Ec 3 
360 10 | Q 10 
zo | 8 
0 0 
l 
Soil Depth (mm) Soil Depth (mm) 
80 : 80 _ ; | 
| r, = 0.46, p< 0.05 Cl | r= -0.12, p< 0.05 d | 
| 
| ° ej. wo 60 | oom ce 6 oe @ 
c a | 
a o& 40, 
- 40 | ee @ Gpareee @ eo ec aupe e | Pal > 7 CUED? ER ee eee e | 
Ss no 2 
3 =O a5 | 
R 20 8 Of SE oeoe | 
wo (eo) | 
> 0 | F 0 | Soneeeene = aoe e e 
be om oe = : 
0 40 60 80 100 0 40 60 80 100 
Soil Depth (mm) Soil Depth (mm) 
| = t 60 
ETS —Vv. < . > . 
= 60 r, = 0.19, p< 0.001 e € r, = -0.31, p =0.014 f 
50 
E 50 ! E 
& E 40 
& 40 *. £ 
E |, ! . E 30 
sia i 5 
2 27H, : Q 20} |e , : 
o o 
oS H ° ‘ ra) 1 
S 105 | | ° ye Qo 10) Bs 8 : 
o 0 | | j sos O 95! . 
0 0 40 60 80 10 0 20 40 60 80 
Total Vascular Plant Cover Total Crytogam Cover 
1,=-0.24,p<0.001 | § q@ a 
r. = -0.24, p< 0. 
# 60° | g 500 = FlintHills = h 
+ se fe Collins 
E 50 | = i 
E | * § 400 \ 
© 40; *. ' iS nN \ 
3) E ua \ 
Ea ; ‘ 8 300 / \ . \ 
9° 8 oO / \ J: \ 
8 / \ 7 » 
& 20 8 : : = 200 Lg ir : 
- i] . ry s / vA \ V/ \ Pace 
10 : : 1° 100 ; \ 
8 |i —i—_+__ Yi 
0 . + 4 d 
aap ey spss : 0 eee near See ner anne es eee —_ 
0 20 40 60 80 00 1998 1999 2000 2001 2002 2003 2005 
Total Vegetation Cover Year 
Fic. 5. Scatter plots of G. minimum, soil depth (millimeters), other vegetation types, with ponding Sy rank correlation 


Site, Dade and St. Clair counties, Missouri). t indicates significance only at the large (1.0 m’) plot scale. 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1155 


occurring on sandy heaths and warrensalong coastal England, has similarities to G. mini- 
mum populations in that population dynamics probably occur at very small scales. It 
has been shown that other vegetation types depress V. ciliata populations significantly. 

The coupled effect of all vegetation upon G. minimum populations further exempli- 
fies that the persistence of this taxon may be dependent upon periodic disturbances. There 
was a significant negative correlation with G. minimum numbers at the subplot level (rs 
=-0.24, P< 0.001) with the combined vegetation cover (cryptogams and vascular plants) 
(Fig. 5g and Table 1). 


Colonization Plots 
The three plots that were created in 1995 by removal of dense moss were not colonized by 
G. minimum during the study. Instead they were eventually recolonized by the moss, al- 
though there were several years following establishment when the habitat appeared suit- 
able for G. minimum. The fact that these plots were nearly surrounded by undisturbed 
areas of dense moss may have precluded the water-transport of G. minimumseeds to the 
plots. Krieger et al. (2003) studied the temporal dynamics of ephemeral plant communi- 
ties in seasonal rock pools on Ivorian inselbergs, monolithic outcrops of granites and 
gneisses along the Ivory Coast of Africa. These rock pools or depressions are typically 
several hundred years old and are seasonally wet with local edaphic that may 
be typically xeric with a thin soil layer. They observed that plant communities in sea- 
sonal pools tend to follow the ‘turnover hypothesis’ which states that species that occur 
in discrete subpopulations tend to be at dynamic equilibrium with extinction and colo- 
nization. The turnover hypothesis is a significant part of island biogeography theory 
(MacArthur & Wilson 1967). The authors also noted that the turnover rate of an annual, 
Cyanotis lanata Benth.(Commelinaceae), was higher than other species. Although turn- 
over was not studied here, the fluctuation in G. minimum population numbers from year 
to year may suggest that turnover does occur. The extent to which it may occur needs 
further study. Furthermore, Krieger et al. (2003) also examined physical characteristics 
of depressional pools including area, mean depth, volume, water depth, and precipita- 
tion for these rock pools. It seems that distance between pools limits the ‘rescue effect’, or 
the migration of a BeEWeen poole G. minimum gene flow among and between 
populations and be ascertained to determine if these populations 
are indeed isolated or have metapopulation characteristics 

To effectively establish G. minimum in additional plots, it is possible to move seeds 
into plots following soil disturbance. This has been done successfully at a highway miti- 
gation site in St. Clair County, Missouri (Smith & Gardner 1999) by moving soil in the 
summer from areas where G. minimum had grown during the spring. It would seem pru- 
dent, however, to select establishment sites that are not subject to rapid recolonization by 
competitive plant species or offset recolonization by competitors with sufficient distur- 
bance. The current study casts doubt on the efficacy of relying on an existing G. mini- 
mum seed bank in the created plots or on the rapid, natural dispersal of seeds into the 
newly-available habitat. 


Climatic Effects 

An aspect of this study that needs further work is the relationship between local and 
regional weather conditions and G. minimum populations. A graphical comparison of G. 
minimum populations between Flint Hill Glades and a relocated population, St. Clair 
County, Missouri, (Smith & Gardner 1999) shows a high degree of symmetry among popu 


1156 BRIT.ORG/SIDA 22(2) 


lation numbers among years, indicating that precipitation and or temperature may play 
a key role (Fig. 5h). The differences between populations in particular years (2001 to 2002) 
may indicate that local precipitation and or temperature events may also influence geo- 
graphically distinct populations. Weather data should be collected on each site for most 
accurate correlations because very local drought conditions at presumed critical life stages 
(fall germination of seed and early spring growth) would not be captured by available 
regional weather station measurements. 


Management Implications 

The long-term monitoring of G. minimum at Flint Hill Glades ecosystem provided in- 
sight on effective management of this threatened species. There is a negative association 
of cryptogams and the population of G. minimum as evidenced by the Spearman rank 
correlations (Fig. 5f and Table Ll). Furthermore, G. minimum seems to utilize habitats where 
there is periodic soil disturbance and minimal competition from mat-forming mosses or 
perennial vascular plants. The level of soil disturbance necessary to keep water-carried 
sand moving into bedrock depressions will be difficult to quantify. 

Geocarpon minimum population bers appear at least partially dependent upon 
the amount of soil present (Fig. 5a, b; Table 1) as indicated by the results of the analysis of 
variance and type II regression, but there are other unknown factors that affect G. mini- 
mum numbers. Some level of grazing by cattle may benefit the plant by maintaining ar- 
eas of loose sand with minimal plant competition, but excessive trampling or nutrient 
deposition would be a concern. Although it seems that many of the moss-dominated 
areas of the glade are currently unsuitable for G. minimum, we have little information on 
how abundant or widespread G. minimum was historically on Missouri’s sandstone glades. 
Fire suppression following settlement of the region has nearly eliminated a critical pro- 
cess that may have previously altered the ratio of bare soil to heavily-colonized soil. 

Management recommendations include providing periodic soil disturbances to the 
glade to assure that loose, sandy soil is available for movement by surface water. The peri- 
odic movement and accumulation of sand into shallow bedrock depressions should facili- 
tate the development of suitable G. minimum habitat. Further studies are needed to address 
the frequency and type of disturbance required to achieve this result. More work is needed 
on the autecology and synecology of this taxon and associated species at other G. mini- 
mum suitable habitats. In addition, the physical envi ent needs to be characterized, as 
well as local and regional climate. It is encouraging that G. minimum was more abundant 
in the plots at this study site in 2003 than during any of the previous nine years and that 
the population is capable of rebounding following a year with little seed production. 


ACKNOWLEDGMENTS 


We thank the following persons for their assistance with monitoring: Mike Skinner, Ann 
Wakeman, John Hays, Dorothy Butler, Craig Tenbrink, Indi Braden, Deborah Renek, Matt 
Parker, Teresa Cochran, and Chris Wieberg. We also would like to acknowledge the U.S. 
Army Corps of Engineers, Stockton Project Office for permission to study Geocarpon at 
Stockton Lake. George Yatskievych, Paul McKenzie, Harold W. Keller, and Lori A. Dameron 
provided invaluable comments on a draft of the manuscript and Brad Jacobs and Nadia 
Navarrete-Tindall provided the Spanish translation of the abstract. 


SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1157 
REFERENCES 


Dausenmire, R. 1959. A canopy coverage method of vegetation analysis. NorthW. Sci. 33:43-64. 

Guo, Q., J.H. Brown and TJ. Vatone. 2002. Long-term dynamics of winter and summer annual commu- 
nities in the Chihuahuan Desert. J. Veg. Sci. 13:565-574. 

Kerry, E.L., J.R. Sincuurst, and S. Cook. 2004. Geocarpon minimum (Caryophyllaceae), new to Texas. Sida 
21:1165-1169. 

Kriecer, A., S. Poremeski, and W. BarTHLoTT. 2003. Temporal dynamics of an ephemeral plant community: 
species turnover in seasonal rock pools on Ivorian inselbergs. Pl. Ecol. 167:283-292. 

MacArtHur, RH. and E.O. Witson. 1967. The theory of island biogeography. Princeton University Press, 
Princeton. 

Mackenzie, K.K. 1914. A new genus from Missouri. Torreya 14:67. 

McInnis, N.C., L.M. Smith, and A.B. Pittman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisi- 
ana. Phytologia 75:159-162. 

Missourl Dept. oF ConservaTION. 2006. Missouri Natural Heritage Database. Resource Science Division. 
Jefferson City, Missouri. 

Moraan, S. 1986. A study of a population of G. minimum in Missouri. Unpublished manuscript. Mis- 
souri Dept.of Conservation, Jefferson City. 

Newson, PW. 2005. The terrestrial natural communities of Missouri. Revised Edition. Missouri Natural 
Areas Committee. Jefferson City. 

Nienaser, M.A. 2005. Geocarpon Mackenzie. In: Flora of North America Editorial Committee, eds. Flora 
of North America north of Mexico, Vol.5 Magnoliophyta: Caryophyllidae, part 2.Oxford Univ. Press. 
New York. Pp. 148-149. 

Orzett, S.L.and E.L. Brioces. 1987. Further additions and noteworthy collections in the flora of Arkan- 
sas, with historical, ecological and phytogeographical notes. Phytologia 64:81-164. 

Quinn, G.P. and MJ. Keouct. 2002. Experimental design and data analysis for biologists. Cambridge 
University Press, United Kingdom. 

Smit, T.E. and J.E. Garpner. 1999. Geocarpon successfully relocated at a western Missouri site. Ecol. 
Restoration 17:91-92. 

Sokal, R.R.and F.J.Ronir. 1995. Biometry. 3 Ed. W.H. Freeman & Co. New York. 

Somers, P, L.R. SmitH, PB. Hamet and E.L. Brioces. 1986. Preliminary analyses of plant communities and 
seasonal changes in cedar glades of Middle Tennessee. ASB Bull. 33(4):178-192. 

THurMAN, C.and E.E. Hickey. 1989. Status report on G. minimum Mackenzie. Provided under contract to 
the U.S. Fish and Wildlife Service, Southeast Region, Atlanta, Georgia. 

U.S. Fish NO Wicowire Service. 1987. Endangered and threatened wildlife and plants; determination of 
threatened status for G. minimum (Geocarpon). Fed. Register 52(115):22930-22933. 

U.S. FisH AND Witoure Service. 1993. Recovery plan for G. minimum Mackenzie. Atlanta, Georgia. 

WATKINSON, A.R., R.P. FreckteTon, and L. Forrester. 2000. Population dynamics of Vulpia ciliata: regional, 
patch, and local dynamics. J. Ecol. 88:1012-1029. 

WATKINSON, A.R. 1990. The population dynamics of Vulpia fasciculata: a nine year study. J. Ecol. 78: 
196-209. 

Warkinson, A.R.and AJ. Davy.1985.Population dynamics of salt marsh and sand dune annuals.Vegetatio 
62:487-497. 

Warkinson, A.R.and J.L. Harper. 1978. The demography of a sand dune annual: Vulpia fasciculata. 1.The 
natural regulation of populations. J. Ecol. 66:1 5-33. 

Zar, J.H. 1999. Biostatistical analysis. 4"" Ed. Prentice Hall, NJ. 


1158 BRIT.ORG/SIDA 22(2) 
Book REVIEW 


Ria Loonuizen. 2006.On Chestnuts: The Trees and Their Seeds. (ISBN 1-903019-832-3, pbk.). 
Prospect Books, Allaleigh House, Blackawton, Totnes, Devon, TQ97DL, U.K. (Orders: 
www.oxbowbooks.com, 1-800-791-9354). $17.95, 136 pp., 51/2" x 71/4". 


“Chestnuts roasting by an open fire...” We are all familiar with the Bing Crosby song White Christmas, but few of 

us are familiar with the history and lore of the chestnut. Growing up in North Texas, I did not see a chestnut tree 

until | was an exchange student in Northern Ireland. The huge tree with its gigantic nuts made quite an impres- 

sion when I came upon it in the dark Irish woods. Ria Loohuizen, a translator living in Amsterdam and the au- 

thor of The Elder, has written a lyrical account of the horse ose and the Sweet chestnut, complete aus 
Wee oy 1 i } rifal f -y 


L 


delight. T] ite book is divided i ti he eecistae Aes¢ ulus hippocastanu and cee 


i 


on the European sweet chestnut, Castanea sativa. It is full of interesting facts. For example, the horse chestnut 
may have gotten its common name from the horseshoe-like shape of its leaf scars a 7. genus name Castanea 
is probably derived from the name of a city in Turkey. The sweet chestnut is native to southern Europe, the 
Caucasus, and North Africa. It is believed to have been brought to Europe by the Celts. A significant section of the 
book is devoted to the culinary applications of pes sweet bee ees is fous only nut — nalen levels of 


vitamin C, according to the author. TI ts, pumpkin 
chestnut soup, Roman-style csinnd with i ntils chestnut sie chestnut cookies, and te chest- 
nut poultry stuffings. T] recipe for a hair conditioner made 


from horse chestnut leaves. Other interesting tips abound: Stich as using crushed horse chestnuts to keep insects 

out of potted plants. Recently, the horse chestnut has gained attention as the source of a treatment for chronic 

venous insufficiency (Suter et al. 2006). The American chestnut tree, Cast dentata, has been a beloved fix- 

ture in the forests of the east coast of the United States, but it has been decimated by chestnut blight, a fungal 

disease imported from Asia. Let us hope that this book is not a eulogy, but rather an ode to the remarkable chest- 

nut.—Marissa Oppel, MS, Collections and Research Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 

76102-4060, U.S.A 

REFERENCES 

Suter, A., S. Bommer., and J. RecuNner. 2006. Treatment of patients with venous insufficiency with fresh 
plant horse chestnut seed extract: a review of 5 clinical trials. Adv Ther. 23:179-190. 


SIDA 22(2): 1158. 2006 


MOUANTITATIVE STUDY OF STYLES AND AGEIENES OF 
TERMINAL AND BASAL FLOWERS OF SCHOENOPLECTUS HALLII 
(CYPERACEAE), A RARE PLANT SPECIES OF 
TRANSIENT WETLAND HABITATS 


Marian Smith, Sara Ammann, Nancy Parker — Paige Mettler-Cherry 


Department of Biology ee of Biology 
Southern Illinois University indenwood Universi se 
Edwardsville Illinois 62026, U.S.A. St. Bis Missouri 63301, 
ABSTRACT 


Schoenoplectus hallii (A. Gray) S.G. Smith (Cyperaceae), a rare plant restricted to wetland habitats, is of conser- 


vation concern throughout its range. ee of the species recently has been clarified; however, quantitative 
i 


Because of 


descriptions of achene and flower morpholo | d life history 
its scarcity and the transient nature of ee any ee -scale study of the species will require the recovery 
of achenes from bulk soil samples and the identification and separation of the dimorphic achenes. The objec- 
tives of this study were to eves, gency and photograph Ge two mechs pes to crane we size pane mor- 


phological differences that wi I g f 
vena in styl phology and ach ithin and amcne, 12 populations in four states. ee each 
achene type ifi ly in size among populations land basal achenes 
are statistically ane and the range of sizes within each achene type is ae than has been previously 
reported. Terminal paid are significa smaller in length, width, beak length, mass and surface area than 
basal achenes, and not in surface ridging. Differences in style morphology are distinct: 
terminal flower styles are ae bifid and consistent in shape, while basal flower styles, which are six 
times longer Een me spies are trifid witha wide variety of branching pea Terminal and basal achenes 
can be sey ly from bulk soil ples using ase oc soil sieves. The vis 

| provided in this study will facilitate the collection ot ntification of eae and 


ey achenes of S. hallii from plants, soil and wildlife. 


RESUMEN 


Sch lectus hallii(A. Gray) S.G. Smith (Cyperaceae) ] ingida a habi le] humed 

es paras ser consetvadaie en ieee su area. La taxonomia de la oad se ha devneads recientemente; sin embargo, 
y la morfologia floral son incompletas y falta informacion de su ciclo 

vital. penis a su escasez y ala earnyeleaa transitoria de sus poblaciones, as estudio a ee a de 


especie requerira la ee de ee a de grandes muestras de suelo y u 
identificacion y separacion de | Por lo tanto, los objetivos a este estudio Pein separar, 
identificar y topralae os dos tipos de aquenio: cuantilica el tamano us eHocnGa mo PICeIee que seran 
utilesen | basales; y 1en la morfologia 


del mn y mans del cogeae en y entre 12 poblaciones en cuatro estados. Aun cada tipo de aquenio varia 
significativamente en tamano entre poblaciones, ms diferencias de tamanio entre aquenios terminales y basales 
son significativas estadisticamente, y el rango de tamano para cada tipo d e lo que se habia 
indicado previamente. Los aquenios terminales son sa teaeganicne mas ee en longitud, anchura, 
longitud el pico, masa y area de su superficie que los aquenios basales, y hay diferencias notables en las costillas 
de B supetticie Las enerenclas en ae morfologia del estilo son oe oe So de las flores Se son 


f] 
bifidos ey onstante | basales. que son seis veces 
] +f: 


1 


Fee | ] 7] f] 7 ] rs | iB a ee Le | 
vValLICQGad ae 
} 


precision y de un modo ee a partir de muestras de 


af 
suelo Slee usando una serie de Subse para suelos. Las visuales y cuantitativas aportadas 


en este estudio facilitaran la recoleccion e identificacion de aquenios recarnales y basales de S. halliia partir de 


plantas, suelo y de la naturaleza. 


SIDA 22(2): 1159- 1173. 2006 


1160 BRIT.ORG/SIDA 22(2) 


INTRODUCTION 


Schoenoplectus hallii (A. Gray) S.G. Smith is a member of the family Cyperaceae. Asa 
Gray (1863) described the species as Scirpus hallii A. Gray, based on specimens collected 
in Illinois. Recent work by Smith (1995) and Smith and Yatskievych (1996), however, has 
emphasized the breakup of the polymorphic supergenus Scirpus sensu lato and resulted 
in the acceptance of Schoenoplectus hallii as the appropriate binomial for the species 
(Smith 1995, 2002). 

The known distribution of S. hallii prior to 1973 extended from Massachusetts to 
Wisconsin and lowa on its northern boundary, westward to Kansas and Nebraska, and 
south to Georgia. The number of states in which populations existed declined from nine 
in 1973 to six in 1997 (McKenzie 1998). It has apparently been extirpated from Massachu- 
setts and Georgia, and is now restricted to the Midwest. In 2000, S. hallii was reported 
from 14 sites in Oklahoma (Magrath 2002); however, the conservation status of the spe- 
cies at these sites is unclear due to the recent discovery of putative hybrids (S. hallii ~ S. 
saximontanus) at some of the sites (Smith et al. 2004). In 2002, surveys for S. hallii re- 
vealed new populations in several states, including Indiana (M. Homoya, pers. comm.) 
and Missouri (pers. obs.) and Texas (O’Kennon and McLemore 2004). In years not suit- 
able for germination, the species may persist only in the seed bank (McKenzie 1998); there- 
fore, it is difficult to obtain accurate estimates of the mE! of wae populations in 
any given year. Although population number an gra tion are not clearly 
defined, S. hallii is considered to be of conservation concern in every state in which it 
occurs (Beatty et al. 2004). 

Schoenoplectus hallii is restricted to wetland habitats (Swink & Wilhelm 1994; 
McClain et al. 1997) in areas characterized by fluctuating water levels (Ostlie 1990; Ostlie 
& Gottlieb 1992; Robertson et al. 1994). The species is thought to have a persistent seed 
bank (Ostlie 1990; Ostlie & Gottlieb 1992; Robertson et al. 1994) that may contribute to 
its potential for population regeneration. Achenes germinate sporadically, depending on 
the availability of wet, exposed habitat (McKenzie 1998), resulting in wide fluctuations 
in population number and size from year to year (Chester 1988; Robertson et al. 1994, 
McClain et al. 1997). It has been suggested that because of loss of suitable wetland habi- 
tat, management of S. hallii sites is necessary to ensure the species’ continued existence 
(Bowles et al. Cee Robertson et al. 1994). 

Scl tus halliiexhibit hicarpy (Bruhl, 1994), as do other members of the 
section Sumpini (ie, S. erectus and S. saximontanus in North America), and achenes pro- 
duced in multi-flowered spikelets near the tip of stems (hereafter designated as “termi- 
nal” achenes or flowers) have been illustrated and/or described by various authors 
(Gleason & Cronquist 1963; Mohlenbrock 1963; Steyermark 1963; Radford et al. 1964; 
Mohlenbrock 1976; Kolstad 1986; Yatskievych 1999; Smith 2002). Although achene size is 
estimated in the taxonomic literature, and varies among authors, a quantitative measure 
of variation in achene size from a representative sample of populations across the range 
of the species is lacking. Basal achenes, which are produced by solitary pistillate flowers 
enclosed within the leaf sheaths at culm bases, are less adequately illustrated and de- 
scribed than terminal achenes. With the exception of a photograph of one basal achene 
(Schuyler 1969) no photographs or illustrations have been published. In addition, no quan- 
titative description of variation in basal achene size within and among populations ex- 
ists. Quantitative data establishing the size and morphology of the basal achenes are im- 
portant for the correct separation and identification of seed bank components and are 


SMITH ET AL., SCHOENOPLECTUS HALLI] ACHENES 1161 


essential for the development of life history studies that estimate the relative reproduc- 
tive contribution of basal achenes to population size. 

In early summer before achenes have been produced, style lobe number is the mor- 
phological character most useful in separating S. hallii from S. saximontanus, because of 
their similar vegetative appearance. As far as we are aware, there has been no large-scale 
study of style size and morphology in S. hallii. Recently, S. hallii and S. saximontanus 
were found growing in mixed populations in Oklahoma (Magrath 2002); therefore, it 
would seem prudent to determine if style lobe number is constant throughout a number 
of widely distributed populations of S. hallii in areas where S. saximontanus is not sym- 
patric to confound the issue. 

The objectives of this study were 1) to develop a simple procedure for separating, 
collecting and identifying achenes from bulk soil samples; 2) to photograph the two 
achene types; 3) to quantify the size and morphological differences that will be useful in 
separating terminal and basal achenes; and 4) to determine the range of variability in 
style morphology and achene size within and among 12 populations in four states (IL, IN, 
KY, and MO) 


MATERIALS AND METHODS 


When possible, living plants were collected in situ and transferred to the greenhouse at 
Southern Illinois University, Edwardsville, IL. Plants were potted in a 75%-sand:25%-pot- 
ting soil mixture in 10 x 10 x 11cm pots, covered with a plastic bag to decrease transpira- 
tional water loss, and placed in flats in 5cm of standing water. The organic content of the 
mixture (%OM = 2.5%) was approximately that of soil at the study sites (mean %OM = 
2.6+0.5%, analyses conducted by Alvey Labs, Belleville, IL). Plants were given identifying 
numbers and mature terminal acl (and basal achenes, when present) were collected 
and saved for measurement. Otherwise, basal achenes were collected as they matured. 
Site names, acronyms, approximate locations (exact locations are not listed to pro- 
tect population sites) and dates of collection are listed in Table 1. At three sites that were 
without plants (SP FO and KY), but were known to have had populations of S. hallii within 
the past 5 years, soil cores were collected to provide a seed source. In all cases, plants or 
cores were collected at regular intervals along transects from across the known extent of 
the population. To ensure accurate identification of achene types, initial collections were 
of terminal and basal achenes that were attached to the parent plant. After achene iden- 
tification was established, others were collected from soil samples. Whenever possible, 
pairs of terminal and basal achenes were selected, using a random number table, from 15 
plants and photographed, measured and weighed in the laboratory. For sites without 
plants, soil sieves (U.S. Standard by Fisher Scientific Co) #16 (1.18 mm mesh), #18 (1.00 
mm) and #20 (0.841 mm) were used to separate basal and terminal achenes. Identifica- 
tion was verified using a dissecting microscope. As some achenes that appeared to be 
healthy and mature fragmented when pressed between the thumb and forefinger, all 
achenes were given this preliminary test before being measured (Baskin et al. 2003). 
Achene photographs were made using a Nikon CoolPix 995 digital camera mounted 
on a Nikon dissecting microscope (Model SMZ800) fitted with a fiber optic ring-light. 
Length, width, aud beak measurements were taken at a magnification of 40X using a 
Leitz comy pe (Model Laborlux S) anda Bausch and Lomb ocular microme- 
ter that was calibrated with a Leitz stage micrometer. Achene beak length was measured 
from the point where tangential lines drawn at the top of the achene and alongside the 


~=— 


1162 BRIT.ORG/SIDA 22(2) 


Tasie 1.Names, acronyms and county and state locations of 12 Schoenoplectus hallii sites and types of samples 
collected for this study. 


Site State County Date and Sample Collected 
Ebken (EB) IL Mason 2000 dist and soil 
Fornoff (FO) IL Mason 2000 Soil only 

Sand Pond (SP) IL Mason 2000 a | 

Wemker (Wk) IL Mason 2000 Plants and soil 
Indiana (IN) IN Daviess 2002 Plants and soil 
Kentucky (KN) KN Christian 2000 Soil only 

Baptist Camp (BC) MO Scott 2000 Plants and soil 
Howell County (HC) MO Howell 2002 Plants and soil 
Petite Isle (Pl) MO Scott 2002 Plants and soil 
Sherer (SH) MO Scott 1999-2001 Plants and soil 
Waterman (WM) MO Scott 2002 Plants and soil 
West Vaco (WV) MO Scott 2002 Plants and soil 


beak intersected (Fig. LD. As beak length varied considerably, but contributed little to 
achene area, achene surface area was calculated as width x (achene length-beak length). 
Mass was determined using a Mettler analytical balance (Model AT 261 Delta Range). To 
illustrate the contrast in size, one pair of terminal and basal achenes were photographed 
with attached styles (Fig. 2A). To illustrate variability in basal and terminal achene mor- 
phology, three of each achene type were photographed in the same field-of-view (Fig. 
2B). Also, three achenes of each type were photographed in cross-section (Fig. 2C) and 
one pair of achenes was photographed showing the achenes in frontal (Fig. 2D) and pro- 
file (Fig. 2E) views. 

Styles were excised from flowers of each type, floated on water and measured. For 
were made on terminal styles from 15 plants from eight 
populations, but for basal styles, which are scarce and difficult to collect intact, measure- 
ments were limited to 15 plants from one population. Others from eight additional popu- 
lations were observed for style morphology, but not measured. To illustrate some of the 
aera in style morphology observed in basal flowers, drawings were made to scale 
(Fig. 3B- 

rae data sets for terminal and basal achenes were combined for analysis, vari- 
ances were significantly different (P<0.001) between achene types for all dimensions; 
therefore, analysis by two-way ANOVA was rejected and data for each achene type were 
analyzed separately by one-way ANOVA. Equal variance was verified using Levene’s Test 
and data were log transformed, when necessary, to ensure normal distribution. Compari- 
sons of mean values for achene characteristics among sites were determined by 
Bonferroni's t-test. Terminal style measurements, for which data could not be success- 
fully transformed, were analyzed by Kruskal- Wallace one-way ANOVA on ranks, followed 
by Tukey’s t-test for comparison of means. Pooled differences in dimensions between 
achene types were determined by t-test, assuming unequal variances. When matched 
pairs of terminal and basal achenes were available within a population, statistical com- 
parisons were made using a paired-samples t-test. All statistical procedures were calcu- 
lated using SPSS 11.4 (SPSS, Inc. 2002) and are in accordance with Sokal and Rohlf (1981). 
Principle ANOVA statistics, means, and P-values) were reported when differences were 
statistically significant. 


statistical analysis 


SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1163 


1mm 


Fic. 1. Method for determining beak length in basal (left) and terminal (right) achenes of Schoenoplectus hallii. Drawn to scale as 
indicated in the figure. 


RESULTS 


Comparisons among populations 

Terminal achenes.—Terminal achenes varied significantly in every dimension (length, 
width, beak length, mass and surface area) among populations (Table 2, Fig. 4+A-E). 
Achenes from HC were significantly larger than those from all other populations, with 
the exception of beak length (Table 3). There was a wide range in terminal achene size 
and mass (Table 4), even with the removal of the extremely large mean values for HC 
achenes, which, when included, raised the upper limit for every character. 

Terminal styles.—All terminal styles examined were bifid, with the exception of the 
one 4-parted style from HC. There was a much wider range in style length compared to 
the range for any achene character (Table 4). There were significant differences in style 
length among populations (Table 2): the styles of the WK population were longer than 
those from any other site and the styles from the SH site significantly longer than all 
except those from WK (P<0.05). There were no significant differences in style lengths 
among the other six sites. 

Basal achenes.—Basal achenes were more varied in shape than terminal achenes (Fig. 
2B); however, there were significant differences among populations for every basal achene 
character measured (Table 5). Unlike the case with terminal achenes, in which those from 
HC were significantly larger than those from all other sites, no single population had 
consistently larger basal achenes (Fig. 5A-E, Table 6). 

Comparisons between terminal and basal achenes and styles.—Pooled means for every 


1164 BRIT.ORG/SIDA 22(2) 


ean 


didi 


Fi. 2A-E. A. Basal (too) and Saeaer ar ; ere iiaiceylee Headlavial view or nasalad 
terminal achenes ofS. hallii; C. cross sectiona d basal ac f S. hallii; D. close-up of terminal (left) and 
basal achenes of 5. hallii; and E. profile view of basal (left) and terminal achenes of S. Aallii. Bars on all photographs are 1 mm in 
length 


SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1165 


mm 


Fic. 3 AD. Representative styles of A. terminal and B-D. basal flowers of Schoenoplectus hallii. 


dimension measured were significantly larger for basal compared to terminal achenes 
(Table 4, P<0.001, df = 289 for each comparison). Within-population comparisons for the 
nine sites for which achene pairs were available indicated that in seven populations (EB, 
WK, IN, BC, SH, WM, WV), basal achenes were significantly larger for all dimensions. At 
HC, area was not significantly different between terminal and basal achenes (P<0.536, df 


1166 BRIT.ORG/SIDA 22(2) 


Tasie 2. Summary ANOVA statistics for terminal achenes and styles of Schoenoplectus hallii from 12 population 
sites 


Character Source of Variation $s df MS F P 

Length Between pops 2457 11 0.223 28.656 <0.001 
Within pops 1.310 168 0.008 

Width Between pops 3.24] 11 0.295 45,384 <0.001 
Within pops 1.091 168 0.006 

Beak length Between pops 0.056 1] 0.005 10.136 <0.001 
Within pops 0.085 168 0.001 

Area Between pops 23.54 1] 2132 61.447 <0.001 
Within pops 5.829 168 0.035 

Mass Between pops 4.092 1] 0.372 26.263 <0.001 
Within pops 2.380 168 0.014 

Style Between pops 24.163 1] 3.452 15.980 <0.001 
Within pops 27.193 168 0.216 


= 11), and at PI, width for terminal and basal achenes did not differ significantly (Ps0.348, 
= 14) 

All terminal achenes examined had prominent transverse ridging extending the 
width of the achene surface as recently reported for S. hallii achenes from Oklahoma 
populations (Magrath 2002). In contrast, transverse ridging was less conspicuous and 
did not extend the entire width of basal achenes, which had obvious vertical ridges. The 
cross-sectional shape of achenes was generally as described by Yatskievych (1999) (“un- 
equally biconvex in cross-section, sometimes slightly concave on 1 side” and “unequally 
3-angled,” for terminal and basal achenes, respectively); however, both achene types oc- 
casionally varied from these descriptions. Some terminal achenes were plano-convex as 
illustrated in Figure 2 C. 

Styles of basal flowers were approximately 6 times longer than those of terminal 
flowers (Fig. 2A, Table +), and exhibited a wide variety of structural morphologies. Styles 
of all terminal flowers examined were bifid (Fig. 3), with the exception of one style from 
HC that had four style lobes (not shown). All styles from basal flowers were trifid with a 
variety of branching patterns, some of which are illustrated in Fig. 3B-D. 


DISCUSSION 


AC. Martin,a U.S. Fish and Wildlife Service biologist stated that when identifying achenes 
encountered in the stomachs of wildlife, his staff found the description of Scirpus achenes 
in the taxonomic literature was unhelpful in separating species (Martin 1943). Although 
he stated that achenes of some species are distinctive enough to present no problem in 
identification, “the difficulty and danger involved in attempting to distinguish some of 
the other species important to wildlife frequently necessitated noncommittal conserva- 
tism.” These troublesome species included what is now known as Schoenoplectus hallii, 
and, although descriptions of its achenes are far more comprehensive now aes in 1943, 
no taxonomic reference adequately describes the variation of achene size and shape ex- 
isting within the species. Our data clarify and illustrate the distinctions between achene 
types and expand the range of achene dimensions cited in the current published litera- 
ture. While several publications (Beetle 1942; Magrath 2002; Schuyler 1969; Smith & 
Yatskievych 1996; Smith 2002) stated clearly that S. hallii exhibits amphicarpy, many 


SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1167 


A. : é 
aia D 3.0 
2.5 
Se 1.5- — 
E 2.0 “Ee 
Ps = 
= 1.0 4 1.5 Pi 
x <x 
1.0 
0.5 4 
0.5 
0.0 0.0 
1.2 
2.0 4 
1.0 
=> 15-4 pret 
= 08 G&D 
£ £ 
— — 
= w 
S 10- Oe <8 
= = 
0.4 
0.5 
0.2 
0.0 0.0 
0.16 4 
5 
0.14 4 
7 0.12 4 4 
= 0.104 = 
<= 
= a £ 
5 0.08 + 2 
<~ 0.06 — 20 
s : 
co 0.04 
4 
0.02 
0.00 


tT syle Tose ay T Lyd aaa 
EB FO SP WK IN KN BC HC PI SHWMWV EB FO SP WK IN KN BC HC PI SC WMWV 
Site 


Site 


£ Ch. J Wes €, 


12 populations, 


Fic. 4 A-F. A. Length, B. width, C. beak length, D. area, and E. mass of | 
dF. style length of inal f] f S. hallii from 8 populati E + SE of 15 measurements. 


did not (Mohlenbrock 1963; Radford et al. 1964; Steyermark 1963; Kolstad 1986; Rolfsmeier 
1995). Of those that recognize variation in floral types, only three provided estimates of 
basal achene size (Schuyler 1969; Yatskievych 1999; Smith 2002), and none presented data 
to quantify differences between terminal and basal achenes, nor any measure of the vari- 
ance in size within achene type across a range of populations. 

Although there is eclee apreenent between our achene-size data and published 
reports, there are notable differences. With the exception of the three cases where size of 
basal achenes was specifically aesed (ie., Schuyler 1969; Yatskievych 1999; Smith 
2002), we assumed that all published estimates referred to terminal achenes. Six authors 
listed the following estimates of length for terminal achenes: 1.3-1.7 mm (Yatskievych 
1999), 1.5-2.0 mm (Mohlenbrock 1976); “up to” 1.5mm (Kolstad 1986); 1.3-1.5 mm (Gleason 
& Cronquist 1963); “less than” 0.4 mm (Robertson et al. 1994); and 1.3-1.7 mm (Smith 
2002). There are three estimates of achene width in the literature: 1 mm (Beetle 1942), 


1168 BRIT.ORG/SIDA 22(2) 


Taste 3. Significant differences in mean length, width, beak length, beak length and mass of terminal achenes 
among 12 populations of Schoenoplectus hallii. All P values calculated by Bonferroni's t-test. 


Length P Beak P Area P Width P Mass P 


HC vs All <0.001 INvs FO  <0.001 HC vs All <0.001 HC vs All <0.001 HC vs All <0.001 

FOvsSP <0.001 vs SP <0.001 FOvs SP <0.001 SHvs SP S0.014 SPvsAll 0.040 

vs SH <0.002. = vs SH <0.001 vs WK <=0.019 SHvsWV S0.018 INvsBC 0.007 
0.002. - - 


INvs SP <0.001 HC vs EB S0.002 vs BC vs PI =0.009 
vs BC <0.006 vsFO <0.001 vs Pl SO001I6 - - vs SH <0.001 
vs SH <0.001 vs SP <0.001 vs SH <0.002 - vs WM <0.001 
vs Pl <0.018 vs WK <0.003 vs WM =0.002 ~ - = 
vs WV <0.011 vs SH 0.001 vs WV 0.001 - - - - 


Taste 4. Pooled means + SE and range of values for length, width, beak length, area and mass for terminal (T) 
and basal (B) achenes, and styles of terminal achenes of Schoenoplectus hallii from 12 population sites. *Omit- 
ting extreme values from Howell County **Mean values of basal styles from SH only. 


T Mean T Range T Range* B Mean Brange 
Length 1.52+0.01 1.14-2.7 1.14-1.76 2.30+0.03 1.20-3.11 
Width 1,32+0.02 0.88-1.88 0.88-1.50 1.43+0.01 1305-1.93 
Beak length 0.11+0.00 0.04-0.22 0.04-0.18 0.34+0.03 0.11—-0.90 
Area 1.88+0.03 1.12-3.32 1.12-2.42 2.81+0.04 1.64-4.32 
Mass 0.60+0.01 0.23-1.13 0.23-0.95 1.68+0.04 0.85-2.83 
Style 3.254+0.06 1.30-6.00 1.30-6.00 17.07+0.89** 12.00-22.00** 


1.2-1.5 mm (Radford et al. 1964) and 1.2-1.3mm (Smith 2002). With the exception of the 
measurement reported by Robertson et al. 1994), which undoubtedly refers to beak 
length, as previously stated in Schuyler (1969), rather than achene length, all measures 
of terminal achenes are within a reasonable range; however, the incomplete dimensions 
given in any single source create uncertainty in identification or confirmation of achene 
identity. As we have demonstrated, size can vary significantly between sites, and in ex- 
ceptional cases (e.g., HC) the variation is so extreme that the achenes of S. hallii could be 
misidentified if size, as described in the existing taxonomic literature, was included in 
the criteria for identification. As it is not customary for authors of taxonomic literature 
to indicate the sample sizes upon which their estimates are made, it is possible that some 
are based ona single achene, although most cite a range in size, which implies more than 
one measurement. In any case, it is time that a clearer delineation of the range of dimen- 
sions in achenes becomes available in the published literature. 

Relying solely on information in the taxonomic literature, it is possible that some- 
one examining the seed bank of S. hallii could fail to recognize basal achenes as belong- 
ing to the species, and thus underestimate the potential population size or the contribu- 
tion of basal achenes to it. Only three authors described basal achene size: Yatskievych 
(1999) states that they are “slightly larger” than terminal achenes, Smith (2002) stated a 
range from 1.7-2.5 mm long and 1.0-1.3 mm wide, and Schuyler (1969) published a photo- 
graph of one terminal and basal achene pair. While the photograph is the most useful 
representation of the appearance of basal achenes currently available in the published 
literature, neither the verbal nor the visual description is adequate to understand the 


SMITH ET AL., SCHOENOPLECTUS HALLIT ACHENES 1169 


Length (mm) 
in 


Width (mm) 


Beak length (mm) 


Area (mm?) 


Mass (mg) 


z 


9 lati Each bar 


Fic. 5 A-E.A. Length, B. width, C. beak length, D.area, and E f basal act f Sct I t 
represents the mean + SE of 15 measurements. 


1170 BRIT.ORG/SIDA 22(2) 


Taste 5.Summary ANOVA statistics for basal achenes of Schoenoplectus hallii from nine population sites. 


Character Source of Variation SS df MS F P 

Length Between groups 2.5755.929 8 0.322 5.537 <0.001 
Within populations 102 0.058 

Width Between groups 0.4141.969 8 0.052 2.680 0.010 
Within populations 102 0.019 

Area Between groups 0.3411.401 8 0.043 3.106 0.004 
Within populations 102 0.014 

Area Between groups 7.246 8 0.906 5.416 <0.001 
Within populations 17.066 102 0.167 

Mass Between groups 8.584 8 1.068 11.204 <0.001 
Within populations 9.727 102 0.096 


Taste 6. Significant differences in mean length, width, mass and beak length of basal achenes among nine 
populations of Schoenoplectus hallii. All P values calculated by Bonferonni's t-test. 


Length P Mass P 

BC vs IN <0.003 BC vs EB <0.001 
Vs HC <0.001 vs IN 0.014 
vs Pl <0.001 vs HC <0.001 
Width Pe vs Pl <0.001 
Pl vs WV =0.009 vs WV <0.001 
Beak P Pl vs SH <0.001 
EB vs HC <0.006 vs WV <0.001 


magnitude and constancy of the differences between achene types, nor the potential of 
using size to separate achenes. Mean size and mass data, including values for the unusu- 
ally wide terminal achenes at the HC population, indicate that basal achenes average 
15x longer, Ll« wider, 1.5~ greater in area and 2.8x heavier than terminal achenes (Table 4). 

Although data in the present study indicated that there was significant variation in 
morphology among basal achenes, they easily can be distinguished from terminal 
achenes, and at any site lacking S.saximontanusor S. erectus they can be identified easily 
to species. Except for the mixed populations in OK and KS, there are no known sites where 
S. hallii co-occurs with the species with which its basal achenes might easily be con- 
fused. Although we processed from 20-100 soil cores from each of the 12 sites to deter- 
mine the potential seed bank for S. hallii, we did not find any mature achenes of any 
other species that resembled S. hallii basal achenes. 

Various authors have noted that transverse or horizontal ridges are present on the 
surface of achenes of S. hallii (Beetle 1942; Gleason & Cronquist 1963; Magrath 2002; 
Radford et al. 1964; Schuyler 1969; Mohlenbrock 1976; Kolstad 1986; Robertson et al. 1994; 
Rolfsmeier 1995; Yatskievych 1999). Our observations confirm the presence of prominent 
horizontal ridges on all terminal achenes examined; however, they are absent or incom- 
plete on basal achenes, which have conspicuous vertical ridges. Although prominent 
horizontal ridges are limited to terminal achenes, this is universally attributed 
to achenes in S. hallii. 

Cross-sectional shape is often used to separate terminal achenes in S. hallii,described 
as “unequally biconvex in cross section” by Yatskievych (1999), from terminal achenes in 


SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1171 


S. saximontanus, which have a distinctly 3-angled shape (Yatskievych 1999). Our study 
corroborates the general regularity of this feature in S. hallii throughout the 12 popula- 
tions studied; however, as is the case with basal achenes, there are exceptions (Fig. 2C). 

Because all published studies of S. hallii are confined to the species’ taxonomy or 
occurrence aan a and rarity or abundance in various regions), which required no 

large ollection, no authors have suggested any methods for separating achenes 
from bulk soil ae The results of our study indicate that it is possible to separate 
terminal and basal achenes from soil samples using a series of soil sieves. Surface litter 
and a large soil fraction can be separated from achenes by using sieves of various sizes, 
depending upon Be Dip of soil and the nature of the litter. For example, if a sample 
ntain ic matter, one can use a #5 sieve to remove litter, a 
#14 sieve to remove coarse maarieles if the soil is sandy, and then sift the soil through 
sieves #16, #18 and #20 to separate achenes by type. The majority of the basal achenes 
will not pass through the #16 sieve, and the few that do are always retained by the #18 
sieve. A small proportion of the terminal achenes remain in the #18 sieve (as was the case 
with many of HC achenes) with the majority of achenes passing through into the #20 
sieve. In our study, none of the achenes passed through the #20 sieve into the next sieve. 
If the soil is predominantly silt or fine loam, all soil passes through the #20, which facili- 
tates the final collection and counting of achenes. If soil is composed of coarse sand, larger 
grains can be filtered out of the sample using the #14 sieve and finer particles of sand 
will pass through the #18 sieve; however, it is impossible to remove all coarse sand from 
achenes using sieves. Final separation of achenes from sand must be done manually. Al- 
though this final process requires some manual separation of achenes and soil using a 
dissecting microscope, the processing of bulk samples of soil for seeds is greatly facili- 
tated using the graduated sieve method described. For future seed bank and germination 
studies requiring large numbers of achenes, this method will be useful. Based on assump- 
tions of a long-lived seed bank, populations of S. hallii are often listed as “extant” if plants 
existed at the site during the previous 25 years (McKenzie 1998); therefore, seed bank stud- 
ies may be essential to the correct assessment of the species’ status. 

Unlike terminal flowers, which have uniformly bifid styles, with the exception of 
the single 4-parted style from HC, basal flowers have trifid styles as shown in Fig. 2A and 
3B-D. Most references to the number of divisions present in S. hallii styles indicate the 
presence of bifid styles (Steyermark 1963; Kolstad 1986; Rolfsmeier 1995; Smith 2002), 
with only Yatskievych (1999) noting “stigmas 2 (rarely 3 in basal spikelets).” Although 
all basal styles had trifid branching, this only became apparent in some cases when the 
styles were floated on a film of water, which allowed shorter style lobes to separate from 
the main branch of the style (Fig. 3-D). Styles of basal flowers, as previously indicated by 
Yatskievych (1999), are longer than those of terminal flowers (Table 4, Figs. 2A and 3A- 
D). Our data indicate that this difference is significant (statistics not shown, see Table 4 
for mean values) with no overlap in size between the two style types. 

Insummary, significant variation in achene size and shape occurs within and among 
populations; identification and separation of terminal and basal achenes can be accom- 
plished rapidly using a series of sieves; transverse ridging is not a regular or prominent 
feature of basal achenes; and in the vast majority of cases, a bifid style is a consistent 
character in the terminal flowers (we noted one exception). All basal styles examined 
were trifid. Data from this study provide visual and quantitative information that will 
facilitate the collection and identification of terminal and basal achenes of S. hallii from 


1172 BRIT.ORG/SIDA 22(2) 


plants, soil and wildlife and contribute to an accurate assessment of the species’ conser- 
vation status. 


ACKNOWLEDGMENTS 


We thank the Missouri Department of Conservation, U.S. Fish and Wildlife Service and 
the Graduate School at Southern Illinois University, Edwardsville, IL, for financial sup- 
port for this study. We are also grateful toa number of colleagues and students who helped 
in making this study possible: Chairo Shaffer, Dan Free, Bethany Meinardii, Josh Knolhoff 
and Rasheda Johnson who helped with achene collection and identification; Xiaowen 
Wang who helped measure achenes; Michael Homoya who provided achenes and plants 
from Indiana, C.C. Baskin, J.M. Baskin and E. W. Chester for collecting soil samples in Ken- 
tucky; reviewers Paul McKenzie, Larry Magrath and Galen Smith for their helpful com- 
ments; and Paul McKenzie for comic relief and help in field collection of plants and 
achenes. 


REFERENCES 


Baskin, C.C,,J.M. Baskin, E.W. CHester, and M. Smit. 2003.Ethylene as a possible cue for seed germination 
of Schoenoplectus hallii (Cyperaceae), a rare summer annual of occasionally flooded sites. Amer. J. 
Bot. 4:620-627. 

Beatty, B.L., W.F. JENNINGS, and R.C. Rawunson. 2004. Schoenoplectus hallii (Gray) $.G.Sm. (Hall's bulrush):a 
technical ee aon assessment. USDA Forest Service, Rocky Mountain Region. http:// 
222.fs.fed.us/r2/projects/sep/a nents/schoenoplectushallii.pdf. 

BeetLe, A.A. 1942. Studies in the genus or L.V. Notes on the section Actaeogeton Reich. Amer. J. 
Bot. 29:653-656. 

Bow es, M.L., M.M. DeMauro, N. Pavotic, and R.D. Hiesert. 1990. Effects of anthropogenic disturbances on 
endangered and threatened plants at the Indiana Dunes National Seashore. Nat. Areas J. 10 
187-200 

BauHt, J.J. 1994. Amphicarpy in the Cyperaceae, with novel variation in the wetland sedge Fleocharis 
caespitosissima Baker. Aust. J. Bot.42:44-448. 

Cuester, E.W. 1988. An unusual habitat type for three rare Kentucky plants. Trans. Kentucky Acad. Sci. 
49:1-102. 

Gieason, H.A.and A. Cronauist. 1963. Manual of the vascular plants of the northeastern United States 
and Canada. D. Van Nostrand Co., Princeton, New Jersey. 

Gray, A. 1863. Manual of botany of the northern United States. 3" edition. New York. 

Koistap, O.A. 1986. Cyperaceae Juss, the sedge family. In: T.M. Barkley and R.L. McGregor, eds. Flora of 
the Great Plains. University Press of Kansas, Lawrence. Pp. 1059-1113. 

Macratn, L. 2002. Schoenoplectus hallii and S. saximontanus: 2000 Wichita Mountain Wildlife Refuge 
Survey. Oklahoma Native Plant Record 2:54-64. 

Martin, A.C. 1943. Achene characters in the genus Scirpus. Amer. Midl. Naturalist 31:250. 

McCiain, WE, R.D.McCiain, and J.E. Esincer. 1997. Flora of temporary sand ponds in Cass and Mason 
Counties, Illinois. Castanea 62:65-73. 

McKenzie, RM. 1998. Hall’s bulrush (Schoenoplectus hallii) status assessment. U.S. Fish and Wildlife Ser- 
vice. Columbia, MO. 

Mon ensrock, R.H. 1963. The Cyperaceae of Illinois: Scirpus. Amer. Midl. Naturalist 70:1-46. 

Montensrock, R. H. 1976. The illustrated flora of Illinois: Sedges—Cyperus to Scleria. Southern Illinois 
University Press, Carbondale. 


SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1173 


O’Kennon, RJ. and C. McLemore. 2004. Schoenoplectus hallii (Cyperaceae), a globally threatened spe- 
cies new for Texas. Sida 21:1201-1204. 

Ostuie, W.R. 1990. The Nature Conservancy's element stewardship abstract for Scirpus hallii (Hall's 
bulrush). The Nature Conservancy, Arlington, Virginia. 

Ostue, W.R. and S. Gortues. 1992. The Nature Conservancy's element global ranking form for Scirpus 
hallii. The Nature Conservancy, Arlington, Virginia. 

Raprorb, A.E.,H.E. AHLes, and C.R. Beit. 1964. Manual of the vascular flora of the Carolinas. University of 
North Carolina Press, Chapel Hill. 

Rosertson, K.R., L.R. PHituipre, and S.M. GEHLHAUSEN. 1994. The current status of Scirpus hallii_A. Gray, Hall’s 
bulrush, in Illinois. Illinois Natural History Survey Technical Report. Champaign. 

Rorsmeier, S.G. 1995. Keys and distributional maps for Nebraska Cyperaceae, Part 1:Bulbostylis, Cyperus 
Dulichium, Eleocharis, Eriophorum, Fimbristylis, Fuirena, Lipocarpha,and Scirpus. Trans. Nebraska Acad. 
Sci. 22:27-42. 

Scuuyter, A.E. 1969. Three new species of Scirpus (Cyperaceae) in the southern United States. Notul. 
Nat. Acad. Nat. Sci. Philadelphia 423:1-12. 

Smith, M., P. McKenzie, P. Mettter-CHerry, and G. SmitH. 2004. A putative hybrid of Schoenoplectus 
saximontanus and S. hallii (Cyperaceae) from Oklahoma. Sida 21:475-479. 

Smitu, S.G, 1995. New combinations in North American Schoenoplectus, Bolboshoenus, Isolepis and 
Trichophorum (Cyperaceae). Novon 5:97-102. 

Smit, S.G. 2002 enoenanicgas In: Flora of North American Editorial Committee, eds. Flora of North 
America, Vol.23.Mag | linidae (in part): Cyperaceae. Oxford University Press, New 
York. Pp. 44-60. 

Smith, S.G. and G. YatskievycH. 1996. Notes on the genus Scirpus sensu lato in Missouri. Rhodora 98: 
168-179. 

Sokat, R.R.and FJ. RoHir. 1981. Biometry. 2nd Edition.W.H. Freeman and Co., San Francisco. 

SPSS, Inc. 2002. SPSS for Windows. SPSS, Inc. 

STEYERMARK, J.A. 1963. Flora of Missouri. The lowa State University Press, Ames. 

Swink, F.and G. WitHetm. 1994. Plants of the Chicago region. 4th Edition. Indiana Academy of Science, 
Indianapolis. 

Yarskievych, G. 1999. Steyermark’s flora of Missouri. Revised edition, Vol. 1. Jefferson City. 


1174 BRIT.ORG/SIDA 22(2) 


Book REVIEWS 


Davip A. SLePER and JOHN M. POEHLMAN. 2006. Breeding Field Crops, Fifth Edition. (ISBN 0- 
8138-2428-1, hbk.). Blackwell Publishing. 2121 State Ave., Ames, [A 50014-8300, US.A. 
and 9600 Garsington Road, Oxford, OX4 a UK. Otc: ed — QIAO, 515-292- 
3348 fax 1-800-862- “6657, www.blacl I] Lcom, blackwell 
publishing.com,). $84.99, 392 pp., 448 b/w illustrations and photographs, 7" x 10". 
This book is intended for beginning students of plant breeding as well as more advanced students and profes- 
sional Plant prEedels The ns oe was published in oak “and since that time it has been one of the most 
One of the most significant wees ay in plant 


popular plant | the world.’ 
Daeeding since edition 4 (1995) has been cea ation - ens ne Be nomics”—using “tools developed by 
molecular biologists such as DNA marker I g p ple son ae genomes . —. of 


our crop plants.” 
Section L: What is Plant Breeding? 
Section II: The Genetic Basis of Ent ‘Breecig 


Section IIL: Tools of the Plant B gcl ber, ion, fertility-regulating mechanisms, 


biotechnology) 
Section IV: Methods in Plant Breeding 


Section V: Germplasm Resources for Breeding Crop Plants 

Sections VI through IX: Applications: Breeding field crops that are self-pollinated or vegetatively propagated; 
Utilizing hybrid and other miscellaneous breeding procedures 

Section X: Maintenance and Seed Production of Improved Cultivars 


—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


Rosin Wuite. 2006. Daphnes: A Practical Guide for Gardeners. (ISBN 0-88192-752-X, 978- 
0-88192-752-8, hbk.). Timber Press Inc, 133 S.W. Second AVES Suite +50, Portland, 
OR 97204-3527, US.A.(Orders: www.timberpress.com berpress.com, 503- 
227-2878, 1-800-327-5680, 503-227-3070 ao $34.95, 232 pp., color photos, 7" x 9". 
The book is written from the author's 30 years of experience in propagating and growing daphnes, both as a 
commercial horticulturalist and as a home gardener (in his “daphnetum”) in southern England. It is arranged in 
seven chapters: History, Taxonomy, and Morphology; Daphne Species; ao Hybrids; Propagation; Cultiva- 


tion; Pests oo“ Diseases; and aoe Use. Nei any he au a White's book is given to discussion of 36 species, each 
with i ti d with subheadings or ed ‘and “botani- 
cal variants’ tandyor’ ‘named clones.” Various cultivars of the two North A aureola L. (na- 


tive to North Africa and Europe) and D. mezereum L. (native to Central Asia and aaa are included. Excellent 
color ees of many of the species and variants < eee any the 

Daphne includes 70 apecies ne ees J. Halda in 2001) mainly found in Europe and temperate and 
subtropical Asia. The shrubs, from prostrate and mat-forming to erect and up to 5 
meters tall. Plomers are 4-petaled and vary it creamy white to pink, purple, lilac, yellow, and green. The au- 
thor notes that “if they are given the right environment and treatment, daphnes are long-lived plants worthy of a 
place in every garden. What is true is that daphnes are unforgiving of mismanagement ... [but] ... To most of us, 
growing daphnes is a challenge which makes success that much sweeter.’—Guy Nesom, Botanical Research Insti- 
tute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A 


SIDA 22(2): 1174. 2006 


ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE 
LANCE ROSIER UNIT OF THE BIG THICKET NATIONAL 
PRESERVE, HARDIN COUNTY, TEXAS 


Larry E. Brown 


Houston Community College 


Barbara R.MacRoberts 
and Michael H. MacRoberts 


Bog Research 


Holman 740 Columbia 
Houston, Texas 77004, U.S.A. Shreveport, Louisiana 71104, U.S.A. 
and an 
Herbarium, see Branch Science Center Herbarium, Museum of Life Sciences 
Westview Drive Louisiana State University in Shreveport 


88 
Houston, Texas 77055, U.S.A. Shreveport, Louisiana, 71115, U.S.A. 


Paul A.Harcombe, 
Warren W. Pruess, |. penile Elsik 


rcs | a 


Department of Ecolo gy 


Stanley D. Jones 


Dp 


| Research Center 
Rice University PO. Box 6717 
Houston, Texas 77005, U.S.A. Bryan, Texas 77805, U.S.A. 


Biology Herbarium 


ABSTRACT 


An annotated, vouchered checklist is provided of the vascular plant taxa of the 10,100 hectare Lance Rosier Unit, 


Big Thicket National Preserve, Hardin County, in southeastern Texas. Documented for the unit are 694 taxa of 
which 71 (10%) are exotics. 


RESUMEN 


1: 1.] ] ee | 


100 hectareas 


de la reserva Lance . Rosier Unit, Big Packet) Nacional ve, del condado de Hardin, en el sureste de Texas. Se 


documentan para la unidad 694 taxones de los que 71 (10 


INTRODUCTION 


This is the fourth part of an inventory, based upon available voucher specimens, of the 
vascular flora of the Big Thicket National Preserve, in southeastern Texas. In three previ- 
ous papers, we provided an annotated checklist of the Hickory Creek, Turkey Creek, and 
Big Sandy Creek units (MacRoberts et al. 2002; Brown et al. 2005; Brown et al. 2006). In 
this paper, we provide an annotated checklist of the Lance Rosier Unit. 


SITE AND METHODS 


The Big Thicket is located mostly within the longleaf pine region of the West Gulf Coastal 
Plain in southeastern Texas (Parks & Cory 1936; McLeod 1971; Harcombe & Marks 1979; 
Marks & Harcombe 1981; Harcombe et al. 1993). The Big Thicket National Preserve con- 
sists of 12 units scattered over seven Texas counties. The units range from 222 to 10,100 
hectares and total about 34,000 hectares (Peacock 1994). The Lance Rosier Unit consists 
of 10,100 hectares and is located in southern Hardin County (Fig. 1). It is relatively flat 
with poor drainage over most of the area. Elevation ranges from about 14 to 30 meters. 
The Little Pine Island Bayou runs through its western and southern sections. The Lance 


SIDA 22(2): 1175— 1189. 2006 


BRIT.ORG/SIDA 22(2) 


1176 
VN I 
\ 
| secon 
o> al 
ALABAMA - COUSHATTA WOODVILLE 5am 
INDIAN RESERVATION _— —f UPPER NECHES RIVER 
CORRIDOR UNIT 
HILLISTER (BS 
& 
WARREN (DB gt 
Y i 


TURKEY 
CREEK UNIT 


MENARD CREEK 
CORRIDOR UNIT 


aan 


NECHES BOTTOM 
& JACK GORE 
BAYGALL UNIT 


KOUNTZE LA 


f 
SILSBEE 
2} py yh EVADALE 


287 LOWER NECHES RIVER 
69 
. LANCE \ | ORRIDOR UNIT 
ROSIER UNIT \ 
LITTLE PINE ISLAND A a aes ag 
BAYOU CORRIDOR UNIT SY celia a 
i cae Co)— 
.. }SOUR LAKE 
Men 
2 BEAUMONT Che 
a ——_— 
ey lee Oe ce Fug, 
IK Soe 
5 


Fic. 1. Location of the Lance Rosier Unit, Big Thicket National Preserve, Texas. 


Rosier Unit was chosen for inclusion in the Big Thicket National Preserve because of its 
wide range of plant communities (Ajilvsgi 1979; Harcombe & Marks 1979; Watson 1979; 
Harcombe & Marks 1981; MacRoberts & MacRoberts 1998). It is dominated by lower- 
slope hardwood pine forest and wetland pine savanna, but also has flatland hardwood 
forest, floodplain hardwood forest, and baygall thicket (Harcombe & Marks 1979; Marks 
S& Harcombe 1981). Mima mounds are common in the wetland pine savanna sections. 
Fire suppression over the past 30 years has resulted in extensive midstory development 
and shading out of herbaceous vegetation. A map of community types can be found in 
Harcombe and Marks (1979), and details regarding soils, topography, and climate can be 
found in Deshotels (1978), Watson (1979, 1982), Harcombe and Marks (1979), and Marks 
and Harcombe (1981). 


BROWN ET AL., FLORA OF PRESERVE 1177 


Like all units of the Big Thicket National Preserve, the Lance Rosier Unit had not 
previously been the subject of a detailed floristic inventory. 

Larry Brown collected about 750 specimens during 15 field days beginning May 23, 
2004 and ending May 14, 2006. 

The MacRoberts’ collected 73 specimens during a study of wetland pine savanna in 
1997 and 1998. Stanley Jones collected 317 specimens during 2002 and 2003. Geraldine 
Watson collected 87 specimens over several years in the late 1970s and early 1980s. Other 
collectors (Sandi Elsik, Warren Pruess, William Carr, Paul Harcombe, Donna Streng, C. 
Liu, and Eric Keith) gathered 198 specimens. A search of the on line TAMU herbarium 
consortium database found some collections at TEX which were annotated and added to 
the list. 

Because our purpose is to produces a net of taxa known to occur on the Lance Rosier 
Unit, a vouchered specimen was con ed to be the only evidence acceptable for inclu- 
sion in the list. In all, more than 1331 herbarium specimens form the data for this report. 
All specimens at Rice University have been annotated by Larry Brown. A complete set of 
these mounted specimens are to be donated to TAES, the Tracy Herbarium, at Texas AGM 
University. Duplicates collections will be distributed to other herbaria. 

In most cases, nomenclature follows Jones et al. (1997) and Kartesz and Meacham 
(1999). For Dichanthelium we followed a pre-publication key in Diggs et al. (2006). Some 
recent literature sources were followed, especially some nomenclature in the published 
Flora of North America volumes. In cases of multiple collections, no more than four are 
listed for each taxon. 


RESULTS 


The following is an annotated list of the Lance Rosier Unit taxa. Abbreviations used are 
given below. 


CL =C. Liu. His specimens are at Rice University. 

DS = Donna Streng. Her collections are at Rice University. 

DW = David Wolf. His one specimen is at TEX. 

EK = Eric Keith. His collections are at Rice University. 

GW = Geraldine Watson. Her specimens are at Rice University. 

LB = Larry Brown. His specimens are at Rice University except for those few at SBSC. 

MM = Barbara and Michael MacRoberts. Their specimens are at TEX, VDB, and BRIT. Also, 
as noted, one or two are at SBSC. 

PH = Paul Harcombe. His collections are at Rice University 

RJ = Ray Jordan. His collections are at TEX. 

SEWP = Sandra Elsik and Warren Pruess. Their specimens are at Rice University. 

SJ = Stanley Jones. His specimens are at Rice University. 

WRC = William Carr. His specimens are at TEX. 


* = exotic 
+ =see note at end of list. 
RGA NTHACERS ACERACEAE 
vata (Walter) Lindau pees (Chapm.) — Acer rubrum L., SISEWP, 15042, 15041; LB 30287 


RWL ong, LB 30165; SEEKWP 3 
Ruellia caroliniensis (J.-.Gmel.) Sate : 30574: SEEKWP 


AGAVACEAE 
Manfreda virginica (L.) Salisb.ex Rose, LB 31083 


3098 
Ruellia humilis Nutt., SJ 14095, 14069; LB 30030 


1178 


ALISMATA 

Sagtoapop Buchenau,LB 29375, 29363, 30041; 
SJ 1375 

Suna ate a (Engelm.) J.G.Sm., LB 3017 

ANACARDIACEAE 

Rhus copallinum L., SJ 13762;LB 29863 

Toxicodendron radicans (L.) Kuntze, LB 29837, SEWP 


ANNONACEAE 
Asimina parviflora 


APIACEAE 

Centella erecta (Lf) Fernald, SJ 17780 
erophyllum tainturieri Hook. var. tainturieri, LB 30407; 
SJSEWP 1 

Cicuta maculata L., LB 30666; SJ 14091 

*Cyclospermum leptophyllum (Pers.) Sprague ex Brit- 
ton &Wilson, LB 30596 

Eryngium integrifolium Walter, LB 30231, 30101,30596 
30697 


— 


Michx.) Dunal, SJ 13806;LB 31181 


fron ae petal Nutt. ex 7. . ae SJ 13906 


| ly d) / 


rticillata Thunb, Lb 
Onpol ator Le a . ae 30016;MM 


' 


Pea capi ae (Michx.) Raf,, LB 29344; SEEKWP 
3094 


Ptilimnium tor 


n (Elliott) Raf, LB 30254 
is L., LB 29915; SJ 14073; GW 3102 


Can; / | 


APOCYNACEAE 
Trachelospermum difforme (Walter) A. Gray, LB 30246 


AQUIFOLIACEAE 

llex ambigua (Michx.) Torr. SJ 14048 

llex coriacea (Pursh) Chapm., LB 30678, 3011 1a; SJ 
13866,13773 


llex decidua Walter, LB 29329; : regi 
ss longipes Chapman, LB 310 
a Soland., LB 29838; ar 
ies ene Aiton, LB 29983; SJ 13759 


ARACEAE 

Arisaema triphyllum (L.) Schott, SJ ] 3843 
ARALIACEAE 

Aralia spinosa L., LB 29854a 


ARECACEAE 

Sabal minor Uacq.) Pers., SJ 14022;LB 30037 
ARISTOLOCHIACEAE 

Aristolochia serpentaria L., GW 156-2 


ASCLEPIADACEAE 
heer es ae ti subsp. longifolia, SJ 13992; 
29379; MM 387 


asclepic perennis aes ad SEEKWP 3104 
Ascle 1S VUOFIE GUL L LB 
Pp g ' 


Matelea gonocarpos omen ‘ace LB 30592, 29862; 


BRIT.ORG/SIDA 22(2) 


ASPLENIACEAE 

lee eal cele (L : ae Sterns & Poggenb., 
LB 29910; SJ 13848, 1 

pene! 


SUE OTIG 


(Lamarck ) RJ. Jansen var. repens 
“alien RJ. ren LB 29919: SJ 14245 

Ageratina altissima (L.) King & H.E. Rob., GW 2006 

Ambrosia artemisiifolia L., LB 30044, 30872 

Ambrosia psi ee ze WP 068; LB 30872, 30891 

Ambrosia trifida L., L 

Amphiachyris Nie ides (DC.) Nutt, GW 3142 

Arnoglossum ovatum (Walter) H.E. Rob., SEEKWP 3115 


13 
Baccharis halimifolia L., GW 3150 
Bidens aristosa oe Britton, LB 30141, 29397; SJ 
14 


Bidens bipinnat ay 3136, 3149 

Bidens Ronieee L.,, LB 30224, 30213:GW 

Boltonia diffusa Elliott, LB 29998, a ee 3652; SJ 
14027 

Chaptalia tomentosa Vent., MM 3733;LB 31058 

Chloracantha spinosa (Benth.) Nesom, GW 3409 

en horri idulum Michx., LB 30391, 30617;SEWP 3052 

iNiu tin (L.) DC., SJ 14088 


con oe (L.) Cronquist var. canadensis, 
SJSEWP 15090 


Conyza canadensis (L.) Cronquist var. glabrata (A. Gray) 
Cronquist, LB 3006 
+Coreopsis gladiata Walter, LB 30229, 30901;:MM 3662; 
901127 
+Echinacea pallida (Nutt.) Nutt. var. sanguinea (Nutt.) 


Gan D.Thomas, SJ 14098 
Ec epost the LB 30158 
Elephan Raeusch., LB 30053 


(opus 
Elephantopus nudatus A. Gray, LB 30239, 30772; GW 
2412 


Elephantopus tomentosus L., LB 30060, 30050; 30049;WP 


a 
Erechtites hieraciifolia (L.) Raf.ex DC., an SJ 13890 
Erigeron philadelphicus L., SJSEWP. 1 
Erigeron procumbens (Houst. ex ai — WP 088 
Erigeron strigosus Muhl. ex Willd. SJ 13819 
Erigeron tenuis Torr.& A. Gray, SEWP 3073 
Eupatorium capillifolium (Small) Lam.,LB 30193;WP 070 
oan aaa folium L.,LB 30735, 30257;RJ s.n. 
is (DC) Torr.& A. Gray, MM 3641 
Fupatorum chil Guene LB ee PH 901146 
torium perfoliatum L., a 30635, — 

mpaonan rotundifolium L.,LB 2936 
+Eupatorium semiserratum cae - 66, 29979:RJs.n. 
Eupatorium serotinum Michx., LB 30142; RJ s.n. 
Eurybia hemispherica (Alexander) Nesom, LB 30120 
Euthamia leptocephala (Torr. & A.Gray) Greene ex Por- 

ter & Britton, LB 29847, 30143; GW 3139 


PRESERVE 


1179 


BROWN ET AL., FLORA OF 


Gaillardia pulchella Foug. var. pulchella, LB 30625, 29854; 

Gamochaeta coarctata (Willd.) Kerguelen, LB 30623, 
30397 

Gamochaeta pensylvanica (Willd.) Cabrera, LB 30601, 


Gamochaeta purpurea (L.) Cabrera, LB 30398, 30590; 
SEWP 3058; SJ 14264 

Helenium amarum (Raf.) H.Rock var.amarum, LB 29887. 
29878; SJSEWP 15048 

Helenium drummondii H.Rock, SJ 13787;MM 3784, 3786 

Helenium flexuosum Raf.,,LB 29957, 30059; SJ 14170;GW 


101 
Helianthus angustifolium L., MM 3735; CL 901125 
Heterotheca subaxillaris (Lam.) Britton & Rusby, LB 
30194, SJ 14240 


Hieracium gronovii L., LB 30720 

*Hypochaeris Hee (Sch. Bip.) Cabrera var. 
albiflora (Ku Cabrera, SJ 13932 

lva angustifolia nue ex DC.,LB 30071 

va annua L.,WP 062; LB 30249, 30249a 

ue oe (A. Gray) K.L. Chambers ex KJ. Kim, LB 


Liatris acidota Er coe nN. Gray, LB 29996; WRC 17233; 
M 4005, SJ 1 
Liatris a ae MM 3648b; LB 30238, 


3023 

Marshallia graminifolia (Walter) Small LE. Watson, LB 
30008; MM 3378; WRC 10838 

Mikania scandens (L.) Willd., LB 30149, 29976 

Pityopsis graminifolia (Michx.) Nutt., LB 30118; WP 059 

+Pluchea baccharis (Mill.) Pruski, LB 29871, 30692; WRC 


17707 
Pluchea camphorata (L.) DC., LB 30183, 30058 
Pluchea foetida (L.) DC., LB 29848; SJ 14511 
Pseudognaphalium obtusifolium (L.) Hillard & Burtt, LB 
30076 


eas ca (Walter) DC., SEEXWP 3074; 
LB 29864 


Pyrhepappus eae (D. Don) DC., GW 2004; LB 


- ia hirta L.,LB 30652 
Rudbeckia texana (Perdue) P. Cox & Urbatsch, LB 30582; 
SEEKWP 3106 


Silphium gracile A. Gray, CL 901124 
Smallanthus uvedalia (L.) Mack. ex Small, LB 31054 


Solidago canadensis L. var scabra Torr. & A. Gray, LB 


30139 
idago odora Aiton var. odora, LB 30255, 30221, WP 


wn 
Qs 


061 
Solidago rugosa Mill.subsp.asper (Aiton) Cronquist, LB 


29988 
Solidago rugosa Mill. subsp. rugosa, LB 30140, 30884 
Solidago sempervirens L. var. mexicana (L.) Fernald,LB 
62 


Solidago tortifolia Elliott, RJ s.n. 


*Soliva sessilis Ruiz & Pavon., SEEKWP 3077; LB 31168 

*4Soliva anthemifolia Jussieu) Sweet, LB 31233 

*Sonchus asper (L.) Hill, SJSEWP 15092;LB 31160 

Symphyotrichum divaricatum (Nutt.) Nesom,GW 1574; 

Symphyotrichum dumosum (L.) Nesom, LB 30235, 

30240; SJ 14257; MM 3648 

Synenyonientin lateriflorum (L A. we S Love, WP 058 

h LB3 


30881, 30880 
Vernonia missurica Raf., LB 29833, 30889 
12 


Xanthium strumarium L., LB 299 

*Youngia japonica (L.) DC., SJ 14021;LB 31049 
AZOLLACEAE 

Azolla caroliniana Willd., LB 30902 
BERBERIDACEA 

Podophyllum pram L., GW 3048 
BETULACEA 


Carpinus ee Walter, SJ 1 
Ostrya virginiana (Mill) K. te 7 on 14, 14173; LB 
30093 


BIGNONIACEAE 

Bignonia capreolata L., LB 29328, 30091; WP 091 

Campsis radicans (L.) Seem. ex Bureau, LB 29832; SJ 
14250 

Catalpa bignonioides Walter, LB 29340 


BLECHNACEAE 
Woodwardia areolata (L.) T. Moore, LB 30777 
Woodwardia virginica (L.) Sm., LB 29396 
BORAGINACEAE 
a procunpel Mill., LB a ae 
Et 1 ell Vey SEW 

g 


BRASSICACEAE 

Cardamine bulbosa (Schreb. Ex Mull) Britton, Sterns, 
oggenb., GW 3049 

*Cardamine debilis D.Don, LB 30292a 

*Cardamine hirsuta L., LB 30292 

Lepidium virginicum L., LB 30603 

BROMELIACEAE 

Tillandsia usneoides (L.) L., LB 29904; SJ 13845 


BUDDLEJACEAE 
Palynremi tm pr rmbens [ea LB 29896 


pb 


BURMANNIACEAE 

Burmannia capitata UU.F.Gmel.) Mart., LB 30107 
CACTACEAE 

Opuntia stricta (Haw.)Haw., GW 1639 
CALLITRICHACEA 

Callitriche heterophylla Pursh, SJSEWP 15076 
Callitriche fineie Nutt., SJSEWP 3107;LB 31228 
CAMPANULACEA 

Lobelia cardinalis L., = 30167 


1180 


Lobelia flaccidifolia Small, LB 30636, 29362; MM 3920 

Lobelia puberula Michx., SJ 14261;LB 30241 

Lobelia reverchonii B.L. Turner, LB 30230; PH 901144 

Triodanis biflora (Ruiz & Pavon) Greene, LB 30604 

Triodanis perfoliata (L.) Nieuwl., GW 3061 

CAPRIFOLIACEAE 

“Lonicera japonica Thunb., SEWP 3065 

Lonicera sempervirens L., SJ 13929 

Sambucus nigra L. subsp. canadensis (L.) R. Bolli, LB 
303 3105 


Viburnum dentatum L., SEEKWP 3078;LB 2986 1a, 29853 
Viburnum nudum L. var. nudum, SJ 13998; LB 29401, 


CARYOPHYLLACEAE 

*Cerastium glomeratum Thuill., SJSEWP 15066 

Sagina decumbens (Elliott) Torr. & A. Gray, LB 30400; 
SEEKWP 3108 


*Stellaria media (L.) Vill., SISEWP. 15063 

CISTACEAE 

+Lechea minorL., LB 30003 

COMMELINACEAE 

Com: / elir IG virgit 1iCa is LB 301 53 

CONVOLVULACEAE 

Dichondra carolinensis le a 30075; SJ 13849 

Nah vulus sericeus = LB 3 
LG 


~ a 
fain 10ea cordatotriloba Dennst. var. cordatotriloba, LB 
4235 


CORNACEAE 

Cornus florida L., SJISEWP 15043;LB 31047 
Cornus foemina Mill. 8 29938; SJ 13758 
Nyssa aquatica L., SJ 1 

Nyssa biflora Walter, : ae 17365 
Nyssa sylvatica Marshall., LB 29336, 29978 


CRASSULACEA 
Penthorum oe L., LB 30671 


CUCURBITACEAE 
Cayaponia quinqueloba (Raf) Shinners, SJ 14515 


CUPRESSACEAE 
Taxodium distichum (L.) L. C. Rich., LB 30207, 29917 
SEEKWP 3099 


CYPERACEAE 
Carex abscondita Mack., SJ 13851 
Carex albicans Willd. ex Spreng. subsp 

JH. Rettig, SEEKWP 3097;LB 30291 
Carex annectens E.P. Bicknell, LB 30382 
Carex basiantha Steud., SJ 13696;LB 30610 
Carex blanda Dewey, SJ 1391 
Carex bulbostylis Mack., LB 31163 
Carex caroliniana Schwein, LB 30612, 30575 
Carex cherokeensis Schwein., SJSEWP. 15049 


= 


lis (Bailey 


BRIT.ORG/SIDA 22(2) 


Carex complanata Torr. & Hook., LB 30381, 29330; SJ 
2, 13864 
Carex crebriflora Wieg., SJ 14553, 13597 
arex crus-corvi Shuttl. ex Kunze, SEEKWP. 3102 
Carex debilis Michx. var. debilis, SJ 13860;LB 31075 
Carex digitalis Willd. var. asymetrica Fernald, SJ 13852 
Carex flaccosperma Dewey, SJ 13803;LB 30388a; SEWP 
3054 


Carex frankii Kunth, LB 30598; SEEKWP 3090 

Carex Ss Rudge, SJ 14177, 14020 

Carex Elliott, CL 901119;SJ 1 

Carex intumescens Rudge, SEEKWP 31 a ae LB 
31077 


Carex joorii Bailey, LB 30781;SJ 14016 

Carex leavenworthii Dewey, SJSEWP. 15079 

Carex longii Mack., SJ 13863; LB 30581, 2934] 

Carex louisianica Bailey, LB 29845 

Carex oxylepis Torr. & Hook. var. oxylepis,, SJ 13917 

Carex retroflexa Muhl. ex Willd., LB 30406, 30602; SJSEWP 
] 5080 


ba AACS 
triatula MUCH I 


Car J] 

ne iene ia eee SJ 13862 

Carex texensis Bailey, LB 30395; SJSEWP 15056 

Carex triangularis Boeck., SJ 13908;LB 30578 

Carex tribuloides Wahlenb. var. sangamonensis Clokey, 
B 30607; SEEKWP 3088 

Carex verrucosa Muh, SJ 14024 


arex vulpil oidea 


Mich + LB 3 30383 
Cladium mariscus J. Pohl subsp jamaicense (Crantz) 
ae LB 30898, 19238b (SBSC), 19421 (SBSC): 


orr. & Hook. ex Torr, LB 29891a 
pene articulatus : LB 30154, SJ 14058 
*Cyperus entrerianus Boeckl., LB cae 
Cyperus erythrorhizos Muhl.,LB 
Cyperus has LB 30262, a 7 14041 
Cyperus ochraceus < Vahl, LB 30885 
tie odoratus LLB 30712 
[ N Steud, SJ 14060, 14260 
opens, Rottb., SJ 14059; GW 3165 
Cyp Steud., SJ 1 a 14076, 14010 
eae retrorsus ae fea 1427 
Cyperus strigosus L., SJ 1 
Cyperus surinamensis sek - 1426 
Cyperus thyriflorus Jungh.,LB 29905; i 14174; SEEKWP 
al s virens _ - J 13896; LB 2933 
orr.) Chapman, LB 30387, 30163 
Eleocharis ae ae Roemer & J.S. Schultes, LB 
30212, 29946 
Eleocharis microcarpa Torr., LB 30654, 30643; SJ 14550; 


IC 
I | 


Eleocharis montana Roem. & Schult., LB 30890 

Eleocharis montevidensis Kunth, LB 29333; SJ 14070 

Eleocharis tuberculosa R.Br., MM 3858;LB 30216, 29961; 
WRC 17230 


BROWN ET AL., FLUAA UE 


PRESERVE 1181 


Fimbristylis aqutumnalis Roem. & Schult. LB 30157, 
30036; GW 3166b 


*Fimbristylis miliacea (L.) Vahl, LB 30157a, 29989 
Fimbristylis puberula Vahl var. puberula Vahl, SJ 14101; 
LB 31182 


Fuirena breviseta ae a LB 30727,29948;MM 3480; 
WRC 1 
Fuirena bushii Kral, iB ne 30010, 19239; SJ 14296 
Isolepis carinata Hook. & Arn.ex Torr., LB 30409 

Kyllinga brevifolia Rottb., LB a 

Kyllinga odorata Vahl, SJ a 

Kyllinga pumila Michx,, L 

Cada caduca ee hi 3479b , 3540 LB 


Rhynchosp ata (L.) H. Pfeiffer, LB 29880, 29417 
SJ 13770 
Rhynchospora corniculata (Lam.) A.Gray, LB 29369 
Rhynchospora divergens Chapm.,LB 294 
Rhynchospora elliottii A.Dietr., LB 19280;MM 3927, 3948; 


Rhynchospora fascicularis (Michx.) Vahl, LB 30217 
Rhynchospora filifolia A. Gray, LB 30658, 30020; MM 
85,3851 


Rhynchospora globularis eu ) Small, var. globularis 
LB 15998; MM 3864, 

Rhynchospora globularis oo Small var.pinetorum 
(Britton & Small ex Small) Gale, SJ 14074 

Rhynchospora ae : ) Vahl, LB 30260; MM 3646; 
WRC 17703a; SJ 14 

jure ee : Gray, MM 3844c; LB 30645, 

1;5J 14613 
mcg sen (Michx,) Vahl, LB 30079, 30073; 
1; SJ 


rhychospo pie (Baldw. ex Elliott) Thomas, SJ 
4077, MM 394 


Rhyne chespera mixta Britton, LB 29895; SJ 14106 
Rhynchospora perplexa Chapm.ex M.A.Curt.LB 30633, 
354 


Rhynchospora plumosa Elliott, LB 30236; WRC 17355; 


MM 3488b; SJ 13936 
Rhynchospora pusilla ee ex M.A. Curtis, LB 30665, 
30024a, 20417;MM 3 
Rhynchospora rariflora lee LB 30630, 29383, 29353; 


MM 3867 
iychoso ee (Gale) Kral, LB 29416, 30647, 
29390; SJ 1408 


Scirpus cyperinus an LB 30696 


Scleria georgiana Core, MM 3386 (SBSC),3488; SJ 13768; 


LB 15999 (SBSC) 


Scleria oligantha Michx., LB 30608, 30576; SEEKWP 3110 


Sclera pauciflora Muhl. ex ae 30015; SJ 13797 

Scleria reticularis Michx., MM 6 

Scleria Hee Michx., ie ae 29355, 30061; 
SEE 111 


CYRILLACEAE 
Cyrilla racemiflora L.,LB 30006; SJ 14089 


DENNSTAEDTIACEAE 

Pteridium aquilinum (L.) Kuhn, WP 090;LB 30045 

DROSERACEAE 

Drosera brevifolia Pursh, SJ 13769; MM 3781 

Drosera capillaris Poir., LB 29418; SJ 14043; MM 3860; 
GW 3064 


DRYOPTERIDACEAE 
Polystichum acrostichoides (Michx.) Schott, LB 30614, 
30056, 29984 


Woodsia obtusa (Spreng.) Torr.subsp obtusa, SJ 14018 


EBENACEAE 

Diospyros virginiana L., LB 29937a;SJ 14051 

ERICACEAE 

eee canescens (Michx.) Sweet, SJSEWP 
15044; LB 31080 

ese viscosum (L.) Torr. SJ 141 

Vaccinium arboreum Marshall, SJ 13785; LB 29953, 


, 14248 


30280 
Vaccinium corymbosum L., LB 30412, 30223; SJ 13799, 
14167 


ERIOCAULACEAE 

Friocaulon decangulare L., LB 29398; SJ 14156; WRC 
7697; 3100 

anceps Morong, LB 29415; SJ 13774 


EUPHORBIACEAE 
Acalypha gracilens A. Gray, GW 3138;LB 30210, 30715 


lachnocaitlon 


7 
Acalypha virginica L., SJ 14165 
*Caperonia palustris (L.) St.-Hil. GW 3138 
Chamaesyce cordifolia (Elliott) Small, SJ 14198 
Chamaesyce humistrata (Engelm.) Small GW 3162 
Chamaesyce maculata (L.) Small, LB 30201, 29935, 


30713 
Chamaesyce nutans (Lag.) Small, GW 3106, 3168; LB 
J 
pit atric NAj 


hx. WP 069; LB 2992 
Croton glanduoss L. var. glandulosus cael -Arg. LB 
307 


oa monanogynus Michx., LB 30886 
a L.,LB 30732a, 30252, 30099 
Phy ni caroliniensis Walter, LB 30896, 30198, GW 


ee urinaria L., LB 29868; GW 3105, 3140 

*Triadica sebifera (L.) Small, SEEKWP 3096 

*Vernicia fordii (Hemsl.) Airy-Shaw, SEEKWP 3109; LB 
30090 


FABACEAE 
*Albizia julibrissin Durazz., SEEKWP 3083; LB 30248, 
051 
Amorpha fruticosa L.,.LB 30618, 29926 
Baptisia bracteata Muhl.ex Elliott var./aevicaulis (A. Gray 
ex Canby) Isley, LB 31085 
Centrosema virginianum (L.) Benth., LB 29851; GW 3145 


1182 


Cercis canadensis L. var. canadensis, SISEWP 15054 
Chamaecrista fasciculata (Michx.) Greene, LB 29849 
Chamaecrista nictitans (L.) Moench, GW 3137 
peo nies lig Mac. Mill., LB 30667 
m (Michx.) DC., LB 30721 
Desm sel aes (Nutt.) DC. GW 3144 
Desmodium paniculatum DC., LB 30185, 30043; SJ 
nee GW 3152 
Erythrina herbacea L.,LB 3 
Galactia volubilis (L.) ce 29860; GW 3148 
aes tricanthos L., GW 2717 
ttidium vesicarium Jacq.) Harper, LB 30716, 29943, 
9925 


Lespedeza virginica (L.) Britton, GW N20 

SUES syecamienys Hook., GW 3074;LB 31052 
puli LB aie laal 308] 

“Medicago Sarees L., LB 3 


& Rose) B.L. Turner, 


MM 3660 
*Melilotus indicus (L (L ) au ue eae me 31048 


LB 29387, 29373; MM 3660; SJ 13779 
Mimosa strigillosa Torr.& A. Gray, te oak 
een Puneet Benth., LB 312 
usifolia (L.) Irw peers LB 29918 
Sesban crummondi faydb) Cory, SEEKWP 3103; LB 


ae iene (Mill.) McVaugh,LB 30710; WP 065 
oe umbellata (Muhl. ex Willd.) Britton, LB 
57;SJ 14086, 14263 
lps pean Nutt., SJ 1 te LB 29999 
reb., SISEWP 15050 
“Trifolium dubia sibihrce. LB 30403 
Trifolium repens L., SEWP 3062; SJSEWP 15096 

“Vicia hirsuta (L.) S.F.Gray, LB 30402, 31053; SEWP 3067 
Vicia ludovici Nutt. subsp. ludoviciana, GW 3073;LB 


3105] 
Vicia minutiflora D. Dietr., SJSEWP. 15091 
ata sativa L.subsp. nigra (L.) Ehrh., SEWP 3066 
0 


PEF LB 30 


Wisteria eee. (L.) Poir., LB 30619 


FAGACEAE 

Castanea pumila (L.) Mill, SEEKWP. 3116; SJ 13784 

Fagus grandifolia ate WP 178;LB 31078 

Quercus alba L., LB 

Quercus falcata ae LB 30584, 29374, 29977; SJ 
13801 

Quercus hemisphaerica W. Bartram ex Willd., LB 30282; 

J 141 3 

Quercus incana W. Bartram, SJ 14082 

Quercus laurifolia Michx., LB 30591, 30174, 29922 

16466 

Quercus lyrata Walter, LB 29928; SJ 13783; GW 514 

Quercus ee a (Ashe) Ashe, rs 31173 

Quercus m ndica Muench., SJ 1379 

Qu ich i Nutt., LB 29840; SJ 13850 


WN 


BRIT.ORG/SIDA 22(2) 


Quercus nigra L., LB 29385, 29331; SJSEWP. 150843 
Quercus pagoda Raf., SJ 13844, 13838, 14116 
Quercus phellos L.,LB 29391, 30025, 29975 
Quercus stellata Wangenh., SJ 13793 

Quercus velutina Lam., SJ 14056 

Quercus virginica Mill., WP 180 


GENTIANACEAE 
*Centaurium floribundum (Benth.) B.L. Rob., LB 30583 
*Centaurium pulchellum (Sw.) Druce, LB 30597, 30094; 


SEEKWP 30 
Sabatia arenicola Greenm., SJ 13870 
Sabatia calycina A. Heller, LB i 
Sabatia campestris Nutt., SJ 1 
Sabatia gentianoides Elliott, ae = 


GERANIACEAE 
ani lini L., SEWP 3075;LB 30285 


GROSSULARIACEAE 
ltea virginica L., SJ 14118, 14023 


HALORAGACEAE 

Myriophyllum pinnatum (Walter) Britton, Sterns, 
Poggenb.,, SJ 13921 

Proserpinaca palustris L,SJ 13913;LB 30644 

Proserpinaca pectinata Lam., LB 30644; GW 3108; MM 
3475; DS 090 


HAMAMELIDACEAE 

Liquidambar styraciflua L.,SJSEWP 15071 

EE ROPNYLLACEAE 

t.ex Choisy, LB 29958; GW 3109 


HYPERICACEAE 
Hypericum crux-andreae (L.) Crantz, LB 29412, 29994; 
SJ 13788; MM 3389 
Hypericum galioides Lam., LB 30728, 29944; S) 14514 
MM 4001 


Hypericum mutilum L., LB 30002; SJ 14160 
Triadenum walteri (J.G.Gmel.) Gleason, LB 30180, 29892, 
30780; GW 3093 


IRIDACEAE 

Sisyrinchium angustifolium Mill. GW 3054; LB 29358; 

LLB s.n. (SBSC) 

Sisyrinchium teas E.P. Bicknell, SJ 13760, 14255; 
MM 387 ie LB ala 


SisvriNCAhIM 


Be) 
ae 
O 


ae LB 31 
Sisyrinchium rosulatum E. P. he ae 3061; SEWP 
3061 


Sisyrinchium sagittiferum E. P. Bicknell, GW 3082 LB 
JUGLANDACEAE 

Carya aquatica (Michx.) ee — 30085 
Carya illinoensis K. Koch, LB 3 

Carya texana Buckley, LB ve i 13970 

Carya tomentosa (Lam.ex Poir.) Nutt., LB 30611,30579 
Juglans nigra L., LB 29913 


BROWN ET AL., FLORA OF THE LANCE ROSIER UNIT, BIG THICKET NATIONAL PRESERVE 1183 


JUNCACEAE 

Juncus acuminatus Michx., SJ 13898, 14012 

Juncus brachycarpus Weigel, LB 29408; 2: 3899, 14099 

Juncus coriaceus Mack.,LB 29932, 29945;SJSEWP 15068 

Juncus dict Elliott, LB 31167 

Juncus diffusissimus Buckley, SJ 13900, 14008 

Juncus effusus L.,LB 30155; SEWP 3071 

sein elli MONEE: SJ 14061; LB 29342, 31180C 
Rostk,, SJ 14002, 14065; LB 29347 

Juncus nodatus Coville, SJ 13901 

Juncus repens Michx., SJ 14013 LB 30641 

Juncus tenuis Willd., LB 30086; SJ 14074a, 13897 

Juncus validus Coville, LB 29334 


LAMIACEAE 

edeoma hispidum Pursh, LB 31166 
Hyptis alata (Raf) Shinners, LB 29885; MM 3484 
lLycopus rubellus Moench, LB ae PH 901159 
Monarda punctata L., LB 2985 
*Perilla frutescens (L.) Britton, i“ 29890; GW 3052 
Prunella vulgaris L.,SJ 13919 

ycnanthemum albescens Torr. & A. Gray, LB 30196, 

29954 


= 


Salvia coccinea Buchoz ex Etlinger, SJ 14259 

Salvia lyrata L.,LB 30054; SJSEWP 15047 

Scutellaria integrifolia L., SJ 13789, 14549, 14052; MM 
3876 


Stachys crenata Raf.., LB 30396; SEWP 3068 


LAURACEAE 


Persea palustris (Raf.) Sarg., LB 30211, 29866, 29381, 


3 
Fy oO foto eee 


(Nutt.) Nees, LB 30222 


LEMNACEAE 
ea punctata (G. Meyer) Les & DJ. Crawford, LB 
1069 


ae brasiliensis Wedd., LB 30878, 070 
Wolffia aH. Karst., LB 310 
Wolfflla eee (Hegelm.) in. is 31068 


LENTIBULARIACEAE 
Pinguicula pumila Michx., SJ 13702; MM 3761 


LILIACEAE 
MELD aurea welsh LB 30028, 29991;SJ 14094;MM 3639 
L. var. canadense, SJ 13765, 13874 

Hymenocallis liriosme (Raf.) Shinners, SEWP 3072 

Hypoxis rigida Chapm.LB 30233, 311 oe 

Hypoxis wrightii (Baker) Brackett, LB 311 

*Narcissus tazetta L., WP 098 

Nothoscordum bivalve (L.) Britton, ete 5046 

*Ophiopogon jaburan Lodd., LB 3 

cic: croceum (Michx.) men SJ 14548; MM 

3783, LB 31057 

*Zephyranthes candida Herb., LB 29911;SE 4581 

LINACEAE 

Linum medium (Planch, Britton var. texanum (Planch.) 
Fernald, LB 303728a, 30657; SJ 14055; MM 3546 


Linum striatum Walter, LB 30653, 30639; SJ 14169: GW 
3075 


LOGANIACEAE 
Gelsemium a (L.) Aiton f,, LB 29386, 30279; 
SJSEWP 15038 
sare pel (JF. Gmel.) Torr. & A. Gray, LB 29881; 
Mi eal sessi sto te F.Gmel.) G. Don, LB 30012; MM 
3483; PH 9 
LYCOPODIACEAE 
een appressa a ) Cranfill, GW 3435; WRC 
7228; MM 3 
LYGODIACEAE 
*Lygodium japonicum (Thunb.ex Murray) Sw., LB 29888, 
29856; SJSEWP 15045 
LYTHRACEA 
ee cathagenes (Jacq.) J.F. Macbr., GW 3076 
a L.,LB 30600 


LU 


faerie ee Push var. lanceolatum (Elliott) Torr. & 
A. Gray, LB 30042; SJ 14270 
Rotala ramosior (L.) ees LB 3016 


MAGNOLI 
Magnoli sen L., LB 29857;SJ 14239 


Magnolia virginiana Ee SJ 1 4038 


MALVACEAE 

“Abutilon theophrasti Medik., GW 3134 

Hibiscus moscheutos L., LB 30173; GW 3094 
Malvastrum coromandelianum (L.) Garcke, GW 3088 
Modiola caroliniana (L.) G. Don, SEWP 3064 

Sida rhombifolia L., LB 30694 

Sida spinosa L., GW 3163 


MELASTOMATAC 
lutea hates - 30663; MM 3388; WRC 17227;SJ 


1, GW 3090 


4054 
rei mariana L., MM 3489; SJ 14155, 14015;GW 3126 


MORACEAE 
Morus rubra L., LB 29939 


MYRICACEAE 
Morella caroliniensis (Mill.) Small, SJ 14117, 14062 
Morella cerifera (L.) Small, LB 30047; WRC 17356 


OLEACEAE 

SH oUanNys virginicus L., WP 087; LB 29335, 31071 
aroliniana M Mill, S 4175 

Fraxinus eee es hese LB 29924 
*Ligustrum sinense Lour, LB 29865; SJ 14068 


ONAGRACEAE 

Gaura lindheimeri Engelm. & A. Gray, LB 30007; WRC 
17701; SJ 14510; DS 124 

Cua ugg Span LB 30146 

a L., LB 29879; = 14509 

Walter, LB 3015 


Froayint 


of 
lL PpAiasicay, { 
os: 


1184 


Ludwigia glandulosa Walter, LB 29987; SJ 14234; GW 
3104 

Ludwigia grandiflora (Michx.) Greuder & Burdet, GW 
3180 


Ludwigia hirtella Raf.;LB 2995 1,29367;SJ 14171;DS 123 
Ludwigia linearis Walter, MM 3538; LB 29959, 29992 
Ludwigia microcarpa Michx., LB 30684 
Ludwigia octovalvis Jacq.) PH. Raven, LB 30725 
Ludwigia palustris (L.) Elliott, LB 30669, 30159 
Ludwigia peploides (Kunth) PH. Raven, LB 3014 
Ludwigia pilosa Walter, LB 30774, 30122a, a CL 
01157 
sabia sulle L. 
Oe iniata Hill, SEWP 3069, SJSEWP on 
Denotieas: speciosa Nutt., LB 30624; SEEKWP 310 


OPHIOGLOSSACEAE 

Botrychium biternatum (Sav.) Underwood, GW 3051 
Opioalessiinn crotalophoroides Walter, LB 31050 
Ophioglossum petiolatum Hook., LB 30394 


ORCHIDACEAE 

Calopogon tuberosus (L.) Britton, le Poggenb., LB 
29406, 29392; SJ 14079; MM 3 

Listera australis Lindl. SJ 13698 

Spiranthes cernua (L.) L.C. Rich., WP 060 

Spiranthes floridana Metre re LB 2041 

+Spiranthes longilabris M 

Spiranthes odorata a a LB 30894 

Spiranthes praecox (Walter) S. Watson, MM 3863; LB 

178 


,LB 30247 


8(SBSC) 


Spiranthes tuberosa Raf., SJ 14093 
Spiranthes vernalis Engelm. & A. Gray, MM 3919 
OSMUNDACEAE 
smunda regalis | 
29360; WP 098 
OXALIDACEAE 
Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner, LB 


var spectabilis (Willd.) A. Gray, LB 


30 3077 
ee debilis Kunth var. oo (DC.) Lourteig, LB 
0250; SEWP 305 
ae i Pursh, (1) ee 15055 


Beate RCEDE 
L., LB 29907; SJSEWP. 16461 
eaaiicie Gee L. in 30613, 30067 


ee 


L., SJ 14272 PINf 

a echinata Mill.,LB 30 

Pinus palustris Mill, LB 2 

Pinus taeda L.,LB 30144, oo SEEKWP 3092 
PLANTAGINACEAE 

*Plantago major L., LB 30289, 30284 

Plantago virginica L., LB 30404, 29882,; SEWP 3060 
PLATANACEAE 

Platanus occidentalis L., LB 29852 


BRIT.ORG/SIDA 22(2) 


POACEAE 
Agrostis hyemalis (Walter) Britton, Sterns, Poggenb., 
SE 3059; SJ 13868 
Andropogon glomeratus (Walter) Britton, Sterns, 
Poggenb., LB 30188; WP 063 
Andropogon virginicus L., LB 30882, 30172; CL 901126 
Anthaenantia texana Kral, LB 30234; WRC 10840; MM 


1 


3 6 
Aristida longispica Poir. var. geniculata (Raf.) Fernald, LB 


Aristida longispica Poir. var. longispica, LB 30236a 
Aristi dee sae oo ) Vasey, LB 30730, 30729;WRC 
2;MM 354 


Aristida purpurascens Poir. var. virgata (Trin.) Allred, LB 
30237, 30228, 30227, 29956 

Arundinaria gigantea (Walter) Muh. eeaen 5072 

Axonopus fissifolius (Raddi) Kuhlm., SJ 14113; MM 
36 LB 30680, 30674 


Axonopus furcatus (Fluggé) A. S. Hitche., LB 29349 PH 
61;5) 14115 

*Briza minor L., SJISEWP 15053 

“Bromus Seis mo LB 30599; SEEKWP 3095 
Cha m (Michx.) Yates, SJ 14252 
Chasmanthium laxum (L.) Yates var. laxum, LB 29839 
Chasmanthium laxum (L.) Yates var. se 

Wipff & S. Jones, LB 30717 

— rugosa (Nutt.) Nash, LB 30736; MM 3721, 


ied 
HETEPOPP EET (P oir. i 


oe n nda (L.) Pers., LB 30621; SEEKWP 3086 

Dichanthelium aci aie (Desv. ex Poir.) Gould & Clark 
var. angustifolium (Elliott) S.L. Hatch, LB 30588, 
31184, 30035; SJ 14459 

Dichanthelium acuminatum (Sw.) Gould & CA. Clark 
var.acuminatum SJ 14556;LB 29357, 30695 

Dichanthelium acuminatum (Sw,) Gould & C.A.Clark var. 
oe (E.L.Rand & Redfield) Gould & CA.Clark, 


ae thea. acumin oe ey ean & CA.Clark var. 
C.A.Clark, LB 30626, 


GUIUCUTTT MING! 1) IOUT 


346. MM 3482. CL 901 136 
Dich oe boscii (Poir.) Gould & Clark, LB 30609, 
29908, 30876 
nee commnutatuin (Schultes) Gould subsp. 
omm LB 29883, 29982: SJSEWP 15058 
pon commutatum (Schultes) Gould subsp. 
equilaterale (Scribner) Freckman & Lelong, LB 
30063 (SBSC) 
Dichanthelium consanguineum (Kunth) Gould & CA. 
Clark LB 30650; MM 3862 
Dichanthelium dichotomum (L.) Gould var. 
dichotomum, LB 30586, 30585,3064 2a, 29897 
Dichanthelium dichotomum (L.) Gould & C.A. Clark 
subsp. microcarpon (Muhl. ex Elliott.) Freckman & 
eee LB 30642, 29897, 29986 
Dichanthelium dichotomum (L.) Gould subsp. nitidus 
(Lam.) Freckman & Lelong,, LB 31162 


BROWN ET AL., F LYUNA VE 


PRESERVE 1185 


Gould 


a 


Dichanthelium ensifolium (Baldw. ex Elliott 
subsp. ensifolium, LB 30089, 29897 

ae ee laxiflorum (Lam.) Gould, LB 29842; 
SJSEWP 15057 

fe ae ovale (Elliott) Gould & C.A. Clark WRC 


17362 

Dichanthelium ovale (Elliott) Gould & CA. Clark subsp. 
villosissimum (Nash) Freckman & Lelong, LB 30660, 
30651;SJ 14554; WRC 17366 

Dichanthelium polyanthes (Schult.) Mohlenbr., LB 
30589, 31319 

Dichanthelium ravenelii (Scribner & Merr.) Gould & CA. 
Clark, LB 30699 

Dichanthelium scabriusculum (Elliott) Gould & CA, 
Clark, LB 30034, 29980, 29348 

Dichanthelium scoparium (Lam.) Gould, LB 30659, 
30261, 29370; SJ 14042 


(Elliott) Gould, LB 29345, 
30004; SJ 14460 

Dichanthelium tenue (Muhl.) Freckman & Lelong, LB 
29377 

Dichanthelium wrightianum (Scribn.) Freckmann, MM 
3385 

Digitaria ciliaris (Retz.) Koel., LB 30226, 30190, 29903, 
30072 

*Digitaria ischaemum (Schreb.) Schreb. LB 30189 

Echinochloa walteri (Pursh) Heller, SJ 14290 

*Fleusine indica (L poe eS 30078 

Elymus virginicus les SJ 

Eragrostis elliottii ed ie 30187;PH 901148 

Fragrostis hypnoides (Lam.) Britton, Sterns, Poggenb,, 
LB 30177 


Fraor 


tis refracta (Muhl.) Scribn., GW 2400; MM 3654 

Eragrostis secundiflora (J.Presl) subsp. Oxylepis (Torr) S. 
Dsoen LB mais: SJ 14075; PH 901143 

le ides (Munro) Hack., LB 30065 

Hordeum pie tien Nutt.,LB 30405; SEEKWP 3085 

Leersia hexandra Sw., LB 30038 

Leersia lenticularis Michx., LB 30166a; SJ 14172 

Leersia virginica Willd., LB 30197; SJ 13856 

*Lolium arundinaceum (Schreb.) S.J. Darbyshire, SJ 
15095 


Melica mutica Walter, SJ 13697 

ees capillaris (Lam,) Trin., Cl 901128;MM 3656; 
LB 3090 

Pears expansa (Poir,) Trin., GW 2408a; MM 3724 
(SBS 


Oplismenus hirtellus (L.) Beauv., LB 30169, 29916 
Panicum anceps Michx., LB 29899; PH 90114 
Panicum brachyanthum Steud.,LB 30187; CL 901156 
Panicum dichotomiflorum Michx., ee 1, 
Panicum Oe Elliott, LB 3 

Pan he Sera eae 


mitomon J 


*Panicum repens L., LB 2 343 
Panicum rigidulum ee ex Nees, LB 30175;PH 901163, 
901167, MM 3732 


Panicum see re ex Trin, LB 19343, 30019; WRC 
17698; MM 3 


Panicum verrucosum nM ies 3666; LB 30184, 30259 

Panicum virgatum L., LB 

*Parapholis nea) G : cee 3865 

*Paspalum m Poir., LB 29884 

Paspalum floridanum Michx., LB 30192, 30123, 30770 

Paspalum laeve Michx., LB 30775, 30171, 29963; MM 
3663 


Paspalum langei (Fourn.) Nash, LB 29902a; SJSEWP 


16460 

Paspalum lividum Trin., LB 30162 

Paspalum minus Fourn., LB 30879, 30640, 30096, 19238 
(SBSC) 

*Paspalum notatum Fluggé, LB 29964, 29405, 19238 
(SBSC) 

Paspalum plicatulum Michx., SJ 14067 

aoe ae a LB 30026; CL 901158; SJ 

05; MM 3 


Psp ee ens LB 29876, 29870a 
pal teud., LB 29942; GW 3023; SEEKWP 


"3093. 
*Poa annua L., SJSEWP 15061 
Poa autumnalis Muhl. ex Elliott, LB 31055 
*Polypogon monspeliensis Muhl. ex Elliott, LB 30606 
Saccharum baldwinii Sprengel. LB 30168, 29931, 30773 
Saccharum giganteum (Walter) Pers., LB 30147, 30892 
Schizachyrium scoparium (Michx.) Nash var. di eu 
(Hack.) Gould, PH 901145; CL 901122; DW s. 
Sorghum halepense (L.) Pers., LB 30594, 29941; eu 
3084 


Sphenopholis obtusata (Michx.) Scribn., SJ 13915; GW 
2246b; LB 31164 

Sporobolus indicus (L.) R.Br, LB 30622, 29909 

Sporobolus st cael LB14907 (SBSC), 14702 
(SBSC); PH 9011 

Steinchisma ea ae Nash, SJ 13892 

Tridens ambiguus (Elliott) J.A.Schultes, WRC 17234; CL 

01120; GW 2406; MM 3544 
Tridens flavus (L.) Hitche. var. flavus, LB 29923;WP 066 
Tridens strictus (Nutt.) Nash,LB 30145,30258;CL 901121 


m dactyloides (L.) L., SJ 14025 
urocta wianvenylla a (Munro ex Wright) R.Webster, GW 
POLYGALACEAE 
Polygala cruciata var. cruciata L., LB 29995; SJ 14111, 
14425, 14293 
| hiaala incarnata ea 
Polygala leptocaulis Torr. . i - LB 30691, 30687; SJ 
14161, 13993 


Polygala mariana Mill, LB 29400, 30027; MM 3726 
Polygala ramosa Elliott, LB 30017; SJ 14158; MM 3877 


POLYGONACEAE 
Brunnichia ovata (Walter) Shinners, LB 29846 


1186 


‘Polygonum cespitosum Blume var. longisetum (De 
oe yor aan LB al i 
jonu Vieisn., LB 30179; GW 3166a 

Beets udroniaawides Michx, SJ 13891 

Polygonum punctatum Elliott, WP 064; LB 30150 
30084a; GW NO2 

Polygonum virginianum L.,LB 30088, 30209 

*Rumex crispus L.,SJ 13859, Ree 

Rumex chrysocarpus Moris, LB 3 

*Rumex pulcher L., LB 30605; eee 3089 


POLYPODIACEAE 

Pleopeltis polypodioides Andrews & Windham 
subsp.michauxiana (Weatherby) Andrews & 
Windham, LB 30084 


eee 
agalli is L., SISEWP 15051 

Ana fea minima (L.) E.H.L. Krause, LB 31 

Samolus valerandi L. subsp. parviflorus a File LB 
29889; SJ 13831 

RANUNCULACEAE 

Clematis crispa L.,LB 30723, 30580; WP 092:SJ 14262 

Ranunculus hispidus Michx.var.nitidus (Chapm.) T. Dun- 
can, SJSEWP 15064 


ie 


—! 


Ranunculus pusillus Poir., SJSEWP 15074 


RHAMNACEAE 
Berchemia scandens (Hill) K.Koch,LB 29874; SEWP 3049 
Rhamnus caroliniana Walter, WP 099 


ROSACEAE 
Crataegus berberifolia Torr. & A. Gray, SJ 14249 
Crataegus brachyacantha Sarg. & Engelm., LB 30205, 
29930; GW 3041 
Crataegus marshallii Eqglest., SJ 14244 
Crataegus opaca Hook. & Arn., SJ 13761; LB 30206, 
9836, 19244 


Crataegus rhe ae hx., LB 30679 
Crataegus viridis L., SJ 14253 

“‘Duchesnea indi os .) Focke, SISEWP 15062 
Photinia pyrifolia (Lam.) K.R.Robertson & J.B.Phipps,LB 

29402; SJSEWP 15039; SJ 13996 

Prunus caroliniana (Mill.) eh SJ 13909 
Prunus serotina pie SJ 14 08 
“Pyracantha } rita Rehder, SEWP 3055 
“Rosa bracteata J.C.Wendl., SJ 14063 

Rubus argutus Link, LB 30873, ee SEEKWP 3082 
i trivialis Michx., SJISEWP 15052 

ea cantoniensis Lour., LB 30398a 


RUBIACEA 

one sree 5 - SJ 14258;LB 30384 

Diodia teres Walte 

Diodia virginiana L,, ce 2300 sas 3475 

Galium aparine L., SJSEWP 1 

Galium obtusum Bigelow, - 

Galium pilosum Aiton, LB co. 14241; SJSEWP 
16462; GW 3066 


=> 


BRIT.ORG/SIDA 22(2) 


Galium tinctorium (L.) Scop., SJSEWP 15073 
Gardenia augusta Merr., SJ 14143 
Houstonia pusilla Schoepf, LB 30393: aa 15060 
Mitchella repens L., LB 29841, 30074; SJ 13 
Olden landi a boscii Oe Eu LB en on 


*Sherardia arvensis L., ae oe SJSEWP 15082 
Stenaria nigricans (Lam.) Terrell, SJ) 14039; PH 901162; 
31226 


RUTACEAE 
*Poncirus trifoliata (L.) Raf.,SJ 14254 


SALICACEAE 
Salix nigra Marshall., LB 29949 


SARRACENIACEAE 
+Sarracenia alata Wood, LB 30726; SJ 13777; WRC 


SAURURACEAE 
Saururus cernuus L., LB 29835; SJ 14168 


CROPHULARIACEA 
ae fasciculata ie Raf.,SJ 13861, 
Agalinis heterophylla (Nutt.) Small ex ae i: ee 
W 3135, 3401 
Agalinis purpurea (L.) Pennell, LB 30875, 30214,30144a; 


=> 


9 
Agalinis viridis (Small) Pennell, LB 30218 
Aureolaria flava (L.) Farw. var. flava LB 29955, 30001; SJ 
14045, 14236 
Bacopa caroliniana (Walter) Robins.LB 30778 
Bacopa monnieri (L.) Pennell, LB 30203, 29886 
Gratiola brevifolia Raf., LB 29364; SJ 14003; GW 3117 
3916 
Grati Pigpiona Michx,, LB ee ha 3127;PH 901155 
Gratiola virg! nian al. SJISEWP J 
*Lindernia crustacea, (L.) F. He | 29900, 30080 
Mecardonia acuminata (Walter) Small,LB 30200, 30100; 
901142 
Mecardonia procumbens (Mill.) Small, SJSEWP. 15089: 


Micranthemum umbrosum (J.G.Gmel.) Blake, LB 30178, 
30164 

Scoparia dulcis L.,LB 30719, 30204, 29901;SJ 14072 

“Verbascum thapsus L., GW 3047 

*Veronica arvensis L.,LB 30401 

Veronica peregrina L., LB 30401a, 30386 

“Veronica polita Fries, LB 30290 


SELAGINELLACEAE 
Selaginella apoda (L.) Spring, SJSEWP 15077; LB 30281 


SMILACACEAE 

Smilax bona-nox L., SEWP 3050, SJ 14066 

Smilax glauca Walter, LB 30182, 30022;5J 14007 
Smilax laurifolia L., SJ 13778; LB 29389, 30113, 30874 
Smilax pumila Walter, LB 30225, 29894; SJ 13904 
Smilax rotundifolia .,LB 29365 


BROWN ET AL., FLORA OF THE LANCE ROSIER UNIT, BIG THICKET NATIONAL PRESERVE 1187 


milax smallii Morong, LB 30040 


Smilax walteri Pursh, GW 2405 


SOE NASERE 


Physi is seat L. are 30399; GW 3 
L., SEEKWP ae. : 29875 
Solanum elaeagnifel um Cav., SJ 14246; GW 3146 


STYRACACEAE 

Styrax americanus Lam.LB 30384a, 30243; SJ 13910 

Styrax grandifolius Aiton, SJ 14005 

SYMPLOCACEAE 

Symplocos tinctoria (L.) Her, LB 29834, 29378, 30095; 
WRC 17360 

TEE IERID A SENE 

hii (Desv.) Morton, LB 29898; SJ 13933 


TYPHACEAE 

Typha latifolia L., LB 31073 

ULMACEAE 

Celtis laevigata Willd., LB 29861 

Planera aquatica J.F. Gmel., LB 30156, 29932; SJSEWP 


Lantana urticioides Heyek, GW 3141 
Phyla nadiflora (L.) Greene, SEEKWP. 3079; LB 29869; SJ 
14097 


*Verbena brasiliensis Vell., LB 30722; SEEKWP 3080 
Verbena halei Small, a se SEEKWP 3114 
Verbena scabra Vahl, LB 29902 


VIOLACEAE 
Viola lanceolata subsp. vittata (Greene) Russell., WRC 
17357; SJ 13699; MM 3762; LB 30296 
iola palmata L., SJSEWP 15059; SJ 13700; GW 30469; 
LB 30294 
Viola sororia Willd. var missouriensis (Greene) McKin- 
ney, GW 3050; LB 31067 


VISACEAE 
Phoradendron serotinum (Raf) M.C. Johnston, $J 14505; 


VITACEAE 


Ampelopsis arb 


p a (L.) Koehne, WP 067;LB 29934 
Pa ened) iInquetolia (L.) Planch., LB 29850 
Vitis aestivalis Miche LB 30724, 30062; SJ 14105 


Vitis cinerea (Engelm) Millard. var. cinerea, LB 29326; SJ 
141 16a 


15067 
Ulmus alata Michx., LB 29920; SJSEWP 15067 Vitis rotundifolia Michx., SEEKWP 3087; LB 30219 
Ulmus americana L., LB 29858, 30288 00 


XYRIDACEAE 


Vien hi. R | 


yris ambigua Bey.ex Kunth,LB 29419, 30116;SJ 14083: 
GW 3446 


Xyris baldwiniana Schult.,LB 29413; SJ 14045a 


URTICACEAE 

Boehmeria cylindrica (L.) Sw., LB 30616, 29891, 30052 
VALERIANACEAE 

Valerianella radiata (L.) Dufr., SJSEWP 15088 


Xyris difformis Chapm. var. difformis, ol 
a geal Xyris supicai L.C. Rich., LB 30218a; SJ 1 
L.,WP 179 Xyris laxifolia Mart.var.iridifolia cae mn LB 30083; 
ream id io (Sweet) Troncoso, SJ 14247; GW 3107a 
EWP 1 Xyris stricta Chapm. var. obscura Kral, LB 30109, 29950, 
29981; MM 3535 
Notes 


Coreopsis gladata.—We are merging C. linifolia with this species following Strother 


Dichanthelium ovale.— WRC 17362 has the papilose-based hairs of subsp. villosissimum 
but the longer (up to 3 mm) spikelets of subsp ovale (refer to Diggs et al. 2006). Thus we 
are unable to assign a subspecies to this collection. 

Echinacea pallida var. sanguinea.—This globally rare (G3) species was once on the Texas 
rare plant list (Pool et al. 2002; Wildlife Diversity Program 2004). It occurs in many east 
Texas counties (Turner et al. 2003). 

Eupatorium semiserratum.—A|though not mapped in Texas by Turner et al. (2003) nor 
by Siripum and Schilling (2006), plants matching the key characters and description in 
Siripum and Schilling (2006)are found in east Texas. 

Lechea minor.—This is the first report of this species for Texas.; details are to be provided 
in a future publication. 

Pluchea baccharis.—An older name for P. rosea. 

Sarracenia alata.—This species was introduced to the Lance Rosier Unit in the 1960s or 


1188 BRIT.ORG/SIDA 22 


2) 


1970s where it has thrived. It occurs naturally and is common in other parts of the Big 
Thicket region. 
Soliva anthemifolia.—T his name replaces S. mutisii in Flora of North America volume 19. 
Spiranthes longilabris.—T his globally rare (G3) species was once on the Texas rare plant 
list (Pool et al. 2002; Wildlife Diversity Program 2004). In Texas, it occurs in wetland pine 
savannas only in Hardin and Newton Counties (Liggio W Liggio 1999). 
Vicia hirsuta.—Jerry Flook (1975) reported this introduced legume from Shelby County 
and Turner et al. (2003) mapped it only in this county. It is a frequent weed near the site 
of the former Teel residence along the road to Little Pine Island Bayou. 

DISCUSSION 
There are 114 families (113 native) and 352 genera GLI native) for the 694 taxa (612 na- 
tive) on the Lance Rosier Unit list. Ten percent of the taxa are non-native. Eight families: 
Poaceae (96 taxa), Cyperaceae (87 taxa), Asteraceae (84 taxa), Fabaceae (38 taxa), 
Scrophulariaceae (19 taxa), Onagraceae (16 taxa), Rosaceae (15 taxa), Euphorbiaceae (15 
taxa), and Rubiaceae (14 taxa), account for 56 % of the taxa collected. The largest genera 
are Carex (31 taxa), Dichanthelium (21 taxa), Rhynchospora (19 taxa), Cyperus (15 taxa), 
Quercus (15 taxa), and Juncus (12 taxa). 

We compared the present list of species for Lance Rosier Unit with a sample of the 
species reported by Turner et al. (2003) in their Atlas of the Vascular Plants of Texas and 
found that only 74% of the species on the Lance Rosier Unit list are reported as occurring 
in Hardin County in that source. This is not surprising considering that, until recently, 
little focused collecting has been undertaken in the Big Thicket region. Basic floristic 
collecting and documenting in the Big Thicket region are still needed. 

While it is the case that our list is incomplete, as are all floras, and a few taxa re- 
ported here may no longer exist on the unit. Most taxa that have grown in the Lance Rosier 
Unit during the past quarter century have been collected (probably 85+ %). We estimate 
that the Lance Rosier Unit has about 620 to 705 native species/taxa. 


ACKNOWLEDGMENTS 
This study was supported in part by a National Park Service Cooperative Agreement No. 
CA 14001004 to Paul Harcombe who also provided the publication funds. Thanks to Tom 
Wendt and Lindsay Woodruff at TEX for the prompt loan of some critical specimens and 
to Eric Keith of Raven Environmental who reviewed the paper. 


REFERENCES 

Avivsal,G. 1979.Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A.& M.Press, 
College Station. 

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, WW. Pruess, LS. Evsik, and D. JOHNSON. 2005. 
Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National 
Preserve, Tyler and Hardin counties, Texas. Sida 21:1807-1827. 

Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, IS. Evsix, and S.D. Jones. 2006. 
Annotated checklist of the vascular flora of the Big Sandy Unit of the Big Thicket National Pre- 
serve, Polk County, Texas. Sida 22:705-723. 

Desuotets, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. U.S.D.A. Soil Conservation 
Service, College Station, Texas. 


BROWN ET AL., FLORA OF PRESERVE 1189 


Dices, Jr., G.M., B.L. Liescome, M.D. Reeo, and RJ. O’Kennon. 2006. Illustrated flora of East Texas. vol 1. Sida 
Bot. Misc. 26:1-1594. 

FLook, JM. 1975. Additions and corrections to the flora of Texas. Sida 6:114. 

Harcomee, PA. and P.L. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Unpub- 
lished report: U.S. Park Service, Santa Fe, New Mexico. 

Harcomee, PA., J.S. Guitzenstein, R.G. Knox, $.L. Orzett, and E.L. Brioces. 1993. Vegetation of the longleaf 
pine region of the West Gulf Coastal Plain. Proc. Ann. Tall Timbers Fire Ecol. Conf. 18:83-103. 

Jones, S.D., J.K. Wiprr, and PM. Montcomery. 1997. Vascular plants of Texas. Univ. Texas Press, Austin. 

Kaatesz, J.T. and C.A. MeacHam. 1999. Synthesis of North American flora. Version 1.0. North Carolina 
Botanical Garden. Chapel Hill. 

Liccio, J.and A.O. Liccio. 1999. Wild orchids of Texas. Univ. Texas Press, Austin. 

MacRoserts, B.R. and M.H. MacRoseats. 1998. Floristics of wetland pine savannas in the Big Thicket 
National Preserve, southeastern Texas. Phytologia 85:40-50. 

MacRosrats, B.R., M.H. MacRoseets, and L.E. Brown. 2002.Annotated checklist of the vascular flora of the 
Hickory Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795. 

Marks, PL.and PA. Harcomer. 1981.Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr 
567-305: 

McLeop, C.A.1971. The Big Thicket forest of east Texas. Texas J. Sci. 23:221-233. 

Parks, H.B.and V.L. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas Agric. Exp. Sta., 
College Station. 

Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A. & M. Press, College Station. 

Poot, J.M., J.R. SincHurst, D.M. Price, and W.R. Carr. 2002. List of the rare plants of Texas. Wildlife Diversity 
program of the Texas Parks and Wildlife Department and the Nature Conservancy, Austin, Texas. 
Unp;ublished January 2000 edition. 

Siripum, K.C. and E.E. ScHitinc. 2006. Eupatorium. In: Flora of North America, vol. 21. Oxford University 
Press. New York, NY. 

StROTHER, J.L. 2006. Coreopsis section Eublepharis. In: Flora of North America, vol.21.Oxford University 
Press. New York, NY. 

Turner, B.L., H.NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 
24: 1-888. 

Warson, G.E.1979.Big Thicket plant ecology:an introduction. Big Thicket Mus. Publ. Ser.,,No.5, Saratoga, 


Texas. 

Watson, G.E.1982.Vegetational survey of Big Thicket National Preserve. Unpublished report.Big Thicket 
National Preserve, Beaumont, Texas. 

Wivowire Diversity Procram. 2004. A list of the rare plants of Texas. Wildlife Diversity Program of Texas 
Parks and Wildlife Department and The Nature Conservancy of Texas. Unpublished report. 


1190 BRIT.ORG/SIDA 22(2) 


Book REVIEW 


Kenbatt R. Lamkey and Micnari Ler. (Eds.). 2006. Plant Breeding: The Arnel R. Hallauer 
International Symposium. (ISBN 0-8138-2824-4, 978-0-8138-2824-4, hbk.). Black- 
well Publishing. 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington 
Road, Oxford, OX4 2DQ UK. (Ondiees: at 292- vane 55 — gia fax 1-800-862- 
6057, wwwhlackwell lcom I hing.com,). $149.99, 
379 pp., numerous illustrations, rel 


A total of 84 authors contribute a collection of 27 chapters, the first “Plant Breeding: Past, Present, and Furure.’ 


From the Preface: * The world of plant breeding has ee ienced dramatic changes during the [last 45 years]. At 


the institutional level, pternauena) centers of cro] nt have emerged and declined, legal and ethical 
issues have become the private Secor (national encarta: federal governments, universi- 
ies) | li ified and placed greater emphasis on basic res to varietal development. Changes 


in ines inucture (e.g., ons season nurseries, service eleborater ies) and tec ee (e.g., computers, machinery, ana- 


of plant breeders to sialuare more germplasm 


in more we ve in more environments ane to identily ge ane pes that exhibit optimal adaptation to the needs of 


s of nature, and the desires of the market. Nascent developments in basic biological and infor- 


> 


coronal sciences, as -exenmplil fied by the gradual annotation of entire genomes and their gene products, have 
provided additional tools and options for various aspects of plant breeding.”—Guy Nesom, Botanical Research In- 


stitute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


Book NOTICE 
M. Nevin Smit. 2006. Native Treasures: Gardening with the Plants of California. (ISBN O- 
520-24425-7, pbk.). University of California Press, California/Princeton Fulfillment 
Services, 1445 Lower Ferry Road, Ewing, NJ 08018, U.S.A. (Orders: www.ucpress.edu/, 
609-883-1759, 609-883-7413 fax). $24.95, 278 pp., color photos, 7" x 10". 


From the author— This is not a ‘how-to’ gardening book, as practical subjects are addressed here in detail, 


and it is certainly not the comprehensive encyclopedia ... Rather, it combines my own personal thoughts, some- 


times maverick opinions, and experience with a woody measure of plainer facts on selected g cup of native 


plants and their culture. The plant genera described here include most of the major ones in cultivation, but also 


some more obscure but garden-worthy groups, thrown in as whim and personal interests dictate.” Smith divides 


his favorites into “trees” (oaks, madrono), “shrubs” (e.g, wild lilac, manzanitas, flannel bushes, giant poppies), 
“two broader genera” (sage and buckwheat), “subshrubs and herbaceous perennials” (e.g., penstemons, iris, lewisias, 
blue eyes, golden eyes), and “bulbs and corms” (lilies, brodiaeas and friends). He ends with “Small Matters—The 


trouble with annuals”: “We forget that Nature's best displays are occasional and unpredictable. We tell ourselves 
| 


that anything seen over such vast areas s [Heer of annual wildflowers] must be easy to grow. And so they are, ina 


well tended t ses pot. Creating ar splay in the open garden is more difficult—as | have learned through 
harsh experier 


Bae book is Saal at least, aimed toward California gardeners, but surely many of the species could be 


in North America. “California gardeners, like their counterparts the world over, can be woefully 


oO 

ignorant of the flora close at hand. They are influenced by a long and hallowed tradition of importing horticul- 
tural treasures from Europe, Asia, and other far-flung regions.” But .... “A fair number of native plants are now 
used eerenan ea ly with exotics of similar features even by g pater whose notion of geographic a te inthe 
local nursery.” Even for non-gardeners, this is a great book with lots of interesting biology, fun to read, even if 
youre not from California—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort a TX 
76102-4068, U.S.A. 


SIDA 22(2): 1190. 2006 


AN UPDATED, ANNOTATED VASCULAR FLORA OF CADDO PARISH, 
LOUISIANA, WITH NOTES ON REGIONAL 
PHYTOGEOGRAPHY AND ECOLOGY 


Barbara R. MacRoberts and Michael H. MacRoberts 


Bog Research, 740 Columbia 
Shreveport, Louisiana 71104, U.S.A. 


and 
Herbarium, Museum of Life Sciences 
Louisiana State University in Shreveport 
Shreveport, Louisiana /1 


al 


Red River Watershed Management Institute 
Louisiana State University in Shreveport 
Shreveport, Louisiana 7 USA. 


Ga 


ABSTRACT 


It has been a quarter century since a checklist of the vascular flora of Caddo Parish, Louisiana, was published. 

Therefore, we revise and update the Caddo Parish checklist, accepting only vouchered species. We found 1405 

species (1168 native, 237 non-native). Additionally, we examine the Caddo Parish flora in terms of its phytogeo- 

graphical affinities. It is closely associated with the eastern United States and particularly with the southeastern 

United States. Caddo Parish appears to be particularly species rich compared to other parishes and counties of 

approwimately the same size across the southeast. This may be: not only because Caddo I Pa oe is abieise put 
| well-collected Caddo Par 


are particularly ie represented and account for a bigh percentage of West Gulf Coastal a endemic and rare 


oy 


species 


Key Worns: Floristic checklist, Caddo Parish, Louisiana flora 

RESUMEN 
Hace ya un cuarto de siglo que se publico el catalogo de la flora vascular de Caddo Parish, Louisiana. Por ello, 
revisamos y ponemos al dia el catalogo de Caddo Parish, aceptando tnicamente especies con pliegos testigo. 


Eaconanie re is (1168 nativas, 237 no nativas). Ademas, examinamos la flora de Caddo Parish en 
Esta muy asociada con el este y particularmente con el sureste de los 


Eotades Unidos. Caddo een merece ser Se auencnenes rico en especies conn Datade: con otros iaseaael de 


aproximadamente el mismo tamano en el sureste | solo porque C 
porque ha sido ae bien muestreado ee las unidad aparecen en LCaddo Parish, estan 
muy bien representac do y aparece un alto porcentae de S} Sy endémicas 
de la Teas costera cel oeste ael golfo. 

INTRODUCTION 


Caddo Parish, Louisiana, is located in the Red River watershed in the center of the West 
Gulf Coastal Plain (Fig. 1). It is bordered on the north by Arkansas, the west by Texas, the 
east by the Red River, and the south by DeSoto Parish. The only boundary that could be 
considered ecologically significant is the Red River; the remainder are political. Approxi- 
mately equidistant from Caddo Parish are blackland prairies to the west, the Ouachita 
Mountains to the north, the Mississippi River and its extensive floodplain to the east, and 
the Gulf of Mexico to the south. Caddo Parish is located in the Oak-Pine-Hickory phyto- 
geographical region of eastern North America (Braun 1950; Skeen et al. 1993; MacRoberts 


SIDA 22(2): 1191— 1219. 2006 


1192 BRIT.ORG/SIDA 22(2) 


ai 
SS AL 
che nae By 
penis cael pzaans 


Fic. 1. Location of Caddo Parish, Louisiana 


S& MacRoberts 2003a). McLaughlin (in press) places it in the Austroriparan floristic 
subprovince, with some characteristics of Comanchian and Illinoian subprovinces. 

The topography of Caddo Parish is relatively simple. The Red River floodplain cov- 
ers about 28 percent of the 2,283 km sq. parish; the remainder is uplands (Fig. 2). The 
{floodplain forms a continuous north-to-south border along the eastern edge of the par- 
ish. The floodplain can be narrow as when the river approachesa bluff, but it is generally 
several kilometers wide. At the edge of the river are natural levees and swales. Floodplain 
elevations range from approximately 60 meters at the northern edge of the parish to about 
+3 meters at the southern edge. The overall north-south slope is very slight. Sediments 
are almost entirely of Red River alluvial origin and are Pleistocene/Holocene (Edwards 
et al. 1980). 

Uplands flank the floodplain at its western edge. This often involves an abrupt el- 
evation transition generally in the range of 10 to 20 meters. Elevations rise to no more 
than 137 meters in rolling low hills and relatively flat terraces. The upland is geologically 
older than the alluvium and generally consists of Tertiary deposits. The geology, soils, 
and climate of Caddo Parish are described by Teague and Wendt (1994) and are a con- 
tinuum of east Texas (Diggs et al. 2006). 


MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH LOUISIANA 1193 


. ARKANSAS C 


f gs (eA | dD Loe L . 1 J if] j ila rears. 4 1 7A\ 
Fig. 2 
I I 


1194 BRIT.ORG/SIDA 22(2) 


Land surveys conducted by the Government Land Office in the 1830s show that in 
presettlement times the uplands were dominated by oak, pine, and hickory and the flood- 
plains by cottonwood, sycamore, willow, pecan, box elder, ash, cypress, and hackberry, 
with essentially no overlap in species between upland and floodplain (MacRoberts & 
MacRoberts 2005). In some parts of the uplands, pine was nearly absent and oak and 
hickory were dominant; in others, pine reached about 50 percent of the canopy. There 
appeared to have been no monospecific pine forest in Caddo Parish. The distribution of 
woody species appears to be the same today as it was 170 years ago (MacRoberts & 
MacRoberts 2005). 

Historically, prairies were scattered across the region in both the uplands and flood- 
plain (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a). The large “Caddo Prai- 
ries” on the floodplain near modern day Gilliam and Hosston no longer exist, and we 
have little or no information about them aside from the brief descriptions Custis and 
Freeman made in 1806 and those in the Government Land Office survey maps and line 
notes (Flores 1984; MacRoberts & MacRoberts 2005). Canebrakes and cedar forests ap- 
parently were common in the early nineteenth century butare entirely gone today (Flores 
1984; MacRoberts et al. 1997). The Red River Raft, an immense logjam long since cleared, 
caused extensive flooding and the creation of “raft lakes,” the water level of which rose 
and fell with the annual fluctuations of the river (Triska 1984). Wallace, Cross, and Caddo 
lakes are impounded remnants of the raft lakes and continue to be dominated Dy cy- 
press. Oxbow lakes are common on the floodplain. 

While the effects of Native Americans on the Caddo Parish landscape are unknown, 
the effects of Euroamericans have been extensive. The river, its floodplain, and adjacent 
upland ecosystems have undergone major modification over the past two centuries. Es- 
sentially the entire landscape has been converted to farms and plantations and, latterly, 
urban sprawl. The once continuous savanna/forest with its numerous plant communi- 
ties covering hundreds of thousands of hectares is now essentially gone with only frag- 
ments of the natural vegetation remaining and few areas, if any, in virgin condition. Bald- 
cypress swamps, black willow riverbank shrublands, and cottonwood forests hang on 
while some floodplain communities such as cedar forests, canebrakes, and prairies have 
vanished entirely. The upland has also been modified, mainly by urban sprawl, logging, 
fire suppression, and clearing for farms. Vast areas of forest were clear-cut in the latter 
part of the nineteenth and early part of the twentieth century and replanted as pine plan- 
tations. 

Caddo Parish is one of the best collected and reported Louisiana parishes and is cer- 
tainly the best collected and reported parish or county in the “Ark-La-Tex” region—north- 
eastern Texas, southwestern Arkansas, and northwestern Louisiana (Smith 1988: Tho- 
mas & Allen 1993-1998; Turner et al. 2003). 

Caddo Parish was first visited and collected by Peter Custis in 1806 (Flores 1984: 
MacRoberts et al. 1997; MacRoberts & MacRoberts 2004b). Josiah Hale collected in the 
parish in the mid-nineteenth century. N.F Peterson from Louisiana State University and 
Reginald WS. Cocks from Tulane University collected in Caddo Parish in the early twen- 
tieth century. FW. Pennell from the Academy of Natural Sciences in Philadelphia worked 
briefly in the area in 1913 when he collected the only known example of Agalinis 
caddoensis (MacRoberts 1978b; Canne-Hilliker & Debrule 1993). EJ. Palmer from Mis- 
souri Botanical Garden and D.S. and HB. Correll (1941), working out of Duke University, 
collected in the area in the late 1930s. Clair A. Brown from Louisiana State University 


— 


LAUUY PARISH, LOUISIANA 1195 


sporadically collected in Caddo Parish through the mid-century. John W. Thieret, then at 
the University of Southwestern Louisiana in Lafayette (now University of Louisiana in 
Lafayette), collected in Caddo Parish in the 1960s. During the early 1970s, Roselie Overby 
(1974) collected the flora of Caddo Parish for her thesis at the University of Louisiana at 
Monroe (formerly Northeast Louisiana University) under the direction of R. Dale Tho- 
mas, who also collected regularly in the parish into the twenty-first century. Upon begin- 
ning the herbarium at Louisiana State University in Shreveport in the early 1970s, D.T. 
MacRoberts collected the parish for the next three decades. His collections and those of 
various students and faculty resulted in numerous contributions to the flora of Louisi- 
ana (MacRoberts 1977a, 1977b, 1978a, 1978b, 1979a, 1979b, 1980a, 1980b, 1980c, 1984a, 
1984b, 1987, 1989; MacRoberts et al. 1997). Allen et al. (2004) collected mainly grasses in 
Caddo Parish through the late twentieth and early twenty-first centuries. Special studies 
have been undertaken such as those of xeric sandylands in northern Caddo Parish 
(MacRoberts & MacRoberts 1995) and the Bickham-Dickson Park/Red River Watershed 
Research Park on the Red River floodplain in Shreveport (MacRoberts & MacRoberts in 
prep.). Other notable collections are those of Larry Raymond from the Walter Jacobs 
Memorial Park, near Blanchard (Thomas et al. in press). The Louisiana Natural Heritage 
Program has been active in documenting the rare flora of Caddo Parish (Louisiana Natu- 
ral Heritage Program 2006; Reid & Faulkner in press). 

The purpose of this paper is to provide an updated, vouchered checklist of the vas- 
cular flora of Caddo Parish, put Caddo Parish in its phytogeographic context, summa- 
rize data on rare plants found in Caddo Parish, and summarize data on regional endemics. 
The construction of the list has been both aided and hampered by precedents. While 
previous lists often provided information on where specimens might be found, they did 
not do so invariably. Although we located vouchers for most reported taxa, many reported 
taxa could not be located, presumably because the sf were misfiled (lost), out on 
loan, initially misidentified and subsequently annotated to another taxon, or were lo- 
cated in an unspecified herbarium. Additionally, some authors reported taxa for which 
no voucher was collected (ie., “sight records”). 


METHODS 


Using published accounts (MacRoberts 1979a, 1984a, 1989; Gandhi & Thomas 1989; Tho- 
mas & Allen 1993-1998: MacRoberts & MacRoberts 1995; Allen et al. 2004; Louisiana 
Natural Heritage Program 2004, 2006; Thomas et al. in press; MacRoberts & MacRoberts 
in prep.), we compiled a checklist Gwith vouchers where possible) of species reported for 
Caddo Parish. We then checked the list against Kartesz and Meacham (1999) and Flora of 
North America (Flora of North America Editorial Committee 1993-2006) to eliminate 
synonyms and to correct nomenclature. We searched LSUS and NLU for vouchers in ad- 
dition to obtaining loans from BRIT, DUKE, LAF LSU, LTU, MO, NLU, NO, and US. 

Some authors included various waifs and garden and horticultural “escapes” (e.g., 
corn[Zea mays L.], and okra [A belmoschus esculentus (L.) Moench]) and we had to make a 
judgement as to their status in the flora. We decided to eliminate almost all of these un- 
less we had good reason to believe that they are propagating themselves in the wild or at 
least persisting. 

Our taxonomic viewpoint favors “lumping” and we do not concern ourselves here 
with taxa below the species level, the reason being that not only are authors inconsistent 
on the number of infraspecific taxa recognized (e.g., compare Smith 1994; Thomas & Allen 


1196 BRIT.ORG/SIDA 22(2) 


1993-1998: Turner et al. 2003) but also on how to divide (and identify) the often subtle 
morphological differences between certain species. Consequently, when comparing flo- 
ristic richness between various geographic areas, our list should be recognized as a “spe- 
cies’ list, not a “taxa” list. We do not include species on the vouchered list that we have not 
inspected except for some Cyperus determined by D. Ferguson (LSU, pers. comm.) and 
two specimens determined by C. Reid (Louisiana Natural Heritage Program, pers. comm.). 

Nomenclature and authorities follow Kartesz and Meacham (1999) in most cases. For 
acaulescent violets, we follow McKinney (1992) and McKinney and Russell (2002). For 
Gamochaeta, we follow Nesom (200, 2004b). 

In order to put Caddo Parish into a phytogeographic context, we randomly selected 
417 native species from our list and, using Kartesz and Meacham (1999) and Thomas and 
Allen (1993-1998), determined their distribution by state or region across North America 
north of Mexico (see MacRoberts & MacRoberts 2003b for method). On the basis of this 
and other phytogeographical information, we developed a thematic map of the floristic 
affinities between Caddo Parish and the remainder of North America. 

Rare plants often indicate rare habitat and vice versa. Using the Louisiana rare plant 
list (Louisiana Natural Heritage Program 2006), we listed all rare plants found in Caddo 
Parish and their primary habitat. We then summarized these data by habitat. Regionally 
endemic plants also indicate unique habitat, as do singularities. Because Caddo Parish is 
in the center of the West Gulf Coastal Plain, we used a list of endemics in that region to 
determine how many occur in Caddo Parish and with which community they are asso- 
ciated (M.H. MacRoberts et al. 2002). Additionally, Louisiana plant species that occur only 
in Caddo Parish were determined using the Louisiana Natural Heritage Program (2006) 
rare plant list and Thomas and Allen (1993-1998). We listed these and their associated 
habitats. 


RESULTS AND DISCUSSION 


We documented 1405 species in Caddo Parish (1168 native and 237 non-native). The 
vouchered checklist of the vascular plants of Caddo Parish is given after the discussion. 
Figure 3 gives the results of the phytogeographic analysis of Caddo Parish species. Table 
I gives Caddo Parish rare plants/plant community associations. Table 2 gives West Gulf 
Coastal Plain endemic species/community associations that occur in Caddo Parish. Table 
3 lists the Louisiana native species found only in Caddo Parish with their associated com- 
munities. 

The figure of 1405 species for Caddo Parish is higher than that for most parishes and 
counties of similar size throughout the southeastern United States (e.g., Nesom & Brown 
1998; Leidolf et al. 2002; Neyland 2002; Marsico 2005). However, Caddo Parish has been 
better collected than most parishes and counties and additional collecting would bring 
up their species counts. 

North America north of Mexico has approximately 19,000 vascular plant species 
(Thorne 1993; Kartesz @ Meacham 1999). Caddo Parish has 7.4 percent of this total but 
constitutes only 0.01 percent of the land area. Likewise, while Caddo Parish constitutes 
only about | percent of the land area of the West Gulf Coastal Plain, it contains about 36 
percent of the West Gulf Coastal Plain flora. 

The Caddo Parish flora is eastern and specifically southeastern. Caddo Parish shares 
99 percent of its flora with Texas, 98 percent with the remainder of Louisiana, 98 percent 
with Arkansas, 91 percent with Mississippi and Alabama, 88 percent with Oklahoma, 50 


MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH, LOUISIANA 1197 


at el 


| ¥ Se. 30-40 
Kf —Nee“apsnt 
Wir KT ene 

MN Gaxrem: 

ae 


percent with the New England states, but only 21 percent with New Mexico. While Caddo 
Parish has 99 percent of its flora in common with Texas, there is a sharp transition zone 
across Texas in which eastern species drop out and are replaced by western species 
(MacRoberts & MacRoberts 2003b). The transition is much less dramatic to the north 
and east. This pattern is also shown in McLaughlin (in press). 

Thirty-six percent of the Louisiana rare species listed by the Louisiana Natural Heri- 
tage Program (2006) for Caddo Parish are essentially specific to one community: xeric 
sandylands (MacRoberts & MacRoberts 1995; B.R. MacRoberts et al. 2002). The next 
ranked community for rare species for Caddo Parish is calcareous prairie with 16 percent 
(MacRoberts et al. 2003) and saline prairie with 14 percent, followed by pine-hardwood 
forest with 1] percent (Larke & Smith 1994). The remaining 23 percent of rare species are 
spread among six plant communities 

Caddo Parish has 45 of the approximately 100 West Gulf Coastal Plain endemic taxa 
(MacRoberts et al. 2002), of which 64 percent are associated with xeric sandylands. The 
next highest communities for endemics are baygalls and calcareous prairies with 9 and 7 
percent, respectively. The remainder of endemics are spread among five communities 

Of the 15 native Louisiana species occurring only in Caddo Parish, 8 occur in xeric 
sandylands, once again pinpointing this habitat/community as unique. 

Since two ities account for the majority of rare and endemic species in Caddo 
Parish, perhaps some comments on these communities is desirable. 

Xeric sandylands are found in eastern Texas, southeastern Oklahoma, southwestern 
Arkansas, and western Louisiana (McBryde 1933; MacRoberts & MacRoberts 1994, 1995, 


BRIT.ORG/SIDA 22(2) 


1198 
Taste 1.Rare plants/community associations for Caddo Parish. 
Community Number of Rare Plants Percentage 
Xeric sandyland 27 36 
Calcareous Prairie 12 16 
Saline Prairie 10 14 
Pine Harwood Forest 8 1] 
Calcareous Forest 5 7 
Baygall 3 4 
Swamp 3 4 
Small Stream Forest 2 3 
Batture Forest ] 1 
Unknown e) 4 
Total 74 100 


Taste 2.Endemic West Gulf Coastal Plain plant species with community associations for Caddo Parish. 


Community Number of Endemic Plants Percentage 
Xeric sandyland 29 64 
Baygall 4 9 
Calcareous Prairie 3 7 
Pine-Hardwood Forest 2 4 
Floodplain/Bottomland 2 4 
Mayhaw Pond 2 4 
Small Stream Forest | 2 
Barrens | 2 
Unknown 1 2 
Total 45 100 


1996, 1997; B.R. MacRoberts et al. 2002; Diggs et al. 2006). They are open to sparsely wooded 
areas that typically occur on terraces or ridges composed of deep sands, generally of 
marine Tertiary origin. Water and air move through these sands creating a dry, desert- 
like environment. Soils, where undisturbed, are of ten cryptogamic. Lichens Cladonia 
spp.) may be common. Quercus incana is common, but stunted Q. stellata and Q. 
margarettiae are also present. Other characteristic species include Shreneie ry id, 
Cnidoscolus texanus, Coreopsis intermedia, oe. ton aneyr am bITa. Er iogonum 
longifolium, E. multifolium, Froelichia floridana, H Matelea 
cynanchoides, Mirabilis albida, Opuntia humifusa, Paronychia drummondii, Pediomelum 
digitatum, P. hypogaeum, Penstemon murrayanus, Phlox drummondii, Polygonella 
americana, Selaginella arenicola, Streptanthus hyacinthoides, Talinum rugospermum, 
Tetragonotheca ludoviciana, Thelesperma filifolium, Tradescantia reverchonii, Yucca 
louisianensis, and Zornia bracteata. Floristic richness is high in this community. Al- 
though widespread, xeric sandylands are not a dominant community across the West 
Gulf Coastal Plain and are generally found in relatively small patches. At the time of 
European settlement, it has been estimated that xeric sandylands accounted for less than 
one percent of western Louisiana (Lester et al. 2005) and, although there are no pub- 
lished figures, probably much less than this in either Oklahoma or Arkansas. In Texas, 


MACROBERTS AND MACROBERTS, VASCULAR FLORA Of LOUISIANA 1199 


Taste 3. Native Louisiana species found only in Caddo Parish with community associations. 


Agalinis caddoensis (community unknown) 
Callirhoe digitata (prairie? 

Canes dno leiaee (oraie 
Carex nd forest, roadside, saline prairie?) 


Coreopsis intermed poe sandyland) 
Dalea ‘phle eoides (xeric sandyland) 
Dasistoma macrophylla (mixed pine-hardwood forest) 
Helianthemum rosmarinifolium (mima mounds in saline prairie) 
Loeflingia squarrosa (xeric sandyland) 
vai ed RAG CNg aS oe sandyland) 
eric sandyland) 
Prunus an (xeric Sa ie 
spermum (xeric sandyland) 
Thelespe filifolium (xeric sandyland) 
Trillium pusillum (baygall) 


a eer | -| 


especially along the Carrizo formation, xeric common, but for the whole 
of this region this y is unlikely to have amounted to more than 5 percent of the 
area. Today, most xeric sandylands have been destroyed. Again figures are not available, 
but probably less than 5 to 10 percent of the original remain in any natural condition. 
Most of this community has been converted to pine plantations, grazing land, and wa- 
termelon patches. In Caddo Parish, it is decidedly rare and is found only in the northern 
part of the parish near Mira, Ida, Rodessa, and Vivian. Over the last twenty years, we have 
watched considerable acreage of this community vanish under cultivation, urban sprawl, 
and habitat deterioration caused by fire suppression (MacRoberts & MacRoberts 1995, 
1998). Generally speaking, therefore, it represents a minor community by area but is an 
important community because of endemism and locally rare species. 

Calcareous prairies were once widespread across the West Gulf Coastal Plain 
(MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a; Lester et al. 2005). Virtually 
none of this community is left today and none in Caddo Parish although prairies were 
once fairly common not only in the uplands, but on the floodplain as well (MacRoberts 
et al. 1997: MacRoberts & MacRoberts 2004b). While prairies were common in central 
Texas (Diggs et al. 1999), far less than one percent of Louisiana (or Caddo Parish) was ever 
prairie land (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a). Today, Louisi- 
ana prairies can be counted only in the tens to hundreds of hectares. 

Another community that deserves mention because of its rareness is saline prairie. 
This community occurs in several places in the West Gulf Coastal Plain (McInnis et al. 
1993; Keith et al. 2004: Lester et al. 2005). It has received some attention because Geocarpon 
minimum, which is listed as federally threatened, occurs in it. In Caddo Parish, there are 
two known saline prairies. One, about 0.5 ha, was discovere d by Overby (1974); the other, 
a prairie of 11 ha, was found in 2005 by C. Reid of the Louisiana Natural Heritage Pro- 
gram (Barron Road Saline Prairie). Overby found Talinum parviflorum in the prairie she 
located. We examined the Overby prairie several times, and relocated Talinum, but the 
last time we visited the site in March 2006, it had nearly been destroyed by all-terrain 
vehicles (ATVs). Barron Road Prairie is also damaged by ATVs. 

Saline prairie is mostly treeless (mima mounds may be present), with many open 
soil areas known as “slicks” with cryptogamic, high sodium soils, and sparse vegetation. 


1200 BRIT.ORG/SIDA 22(2) 


— 


Characteristic plants include Anagallis minima, Claytonia virginica, Crassula aquatica, 
Eragrostis lugens, Evolvulus sericeus, Houstonia micrantha, Houstonia pusilla, Houstonia 
rosed, Krigia occidentalis, Lepuropetalon spathulatum, Plantago pusilla, and Sporobolus 
vaginiflorus. Ten Louisiana and one globally rare species have been located at the Barron 
Road Saline Prairie. These are Cooperia drummondii, Geocarpon minimum, Gratiola flava, 
Helianthemum rosmarinifolium, Lotus unifoliolatus, Minuartia drummondii, Minuartia 
muscorum, Phacelia glabra, Schoenolirion wrightii,and Talinum parviflorum. 

The two surviving plant collections from the 1806 Freeman-Custis’ Red River Expe- 
dition are Eustoma russellianum (Hook.) G. Don and Veronicastrum virginicum (L.) Farw. 
(Flores 1984, MacRoberts &@ MacRoberts 2004b). Eustoma russellianum does not occur 
anywhere in the region today, and Veronicastrum virginicum is very rare in the region 
and has not been found in Caddo or Bossier parishes since Custis’ original collection. We 
have noted this same pattern with subsequent early collectors in the area. Pennell found 
the only species endemic to Caddo Parish—Agalinis caddoensis—near Shreveport in 1913; 
it has not been found since. Hale, Cocks, Peterson, Correll, Palmer, and other early collec- 
tors found species in the area that are either very rare today or entirely missing (e.g., Buchloe 
dactyloides, Dasistoma macrophylla, Gratiola flava, Helianthus occidentalis, Helianthus 
petiolaris, Onosmodium bejariense, Panicum flexile, Prosopis glandulosis, Sophora affinis, 
and Taenidia integerrima). This suggests that in the last two centuries, a great deal of the 
original habitat has been lost and that the flora was once richer in native species than it 
is today. 


~— 


VOUCHERED CHECKLIST OF VASCULAR PLANTS 


The following is a vouchered list of the vascular plants for Caddo Parish. We list one 
voucher for each species. Key to vouchers: DTM = D.T. MacRoberts. Unless otherwise stated, 
his specimens are at LSUS. RDT = R. Dale Thomas. Unless otherwise stated, his speci- 
mens are at NLU.MM = BR. and M.H. MacRoberts. Unless otherwise stated, their speci- 
mens are at LSUS. Raymond = Larry R. Raymond. The herbarium where his specimens 
are located is always given. Hardy = L.M. Hardy. His specimens are at LSUS. Haynes = 
Robert Haynes. His specimens are at LSUS. Barbour = Philip Barbour. His specimens are 
at LSUS unless otherwise stated. Other collectors are given full citations. An * (asterisk) 
indicates a non-native species. 


PTERIDOPHYTES Polystichum acrostichoides (Michx.) Schott; DTM 2197 
Woodsia obtusa (Spreng.) Torr; DTM 835 
faeces 
platyneuron (L.) B.S.P; DTM 829 ee 
AZOLLACEAE m hyemale L.; Barbour 932 
< ; ae ie Clute; DTM 1700 
Azolla caroliniana Willd.; MM 5935 
BLECHNACEAE Doria eae 
: Isoetes melanopoda Gay & Durieu ex Durieu; RDT 89029 
Woodwardi lata (L.) T. Moore; DTM 1546 
LYGODIACEAE 


Sm.; RDT 33699 


lA m fs 7 si te | 
VVYUUCUVV UT CICt VIFGTEIICCH (L.) 


*L ygodium japonicum (Thunb. ex Murr) Sw.; DTM 1090 
eo incdtes eae 


(L.) Kuhn; DTM 898 


DRYOPTERIDACEAE 

Athyrium filix-femina (L.) Roth; DTM 1221 

*Cyrtomium falcatum (L.f.) K.Presl; DTM 113 (question- 
ably naturalized) 

Onoclea sensibilis L.; DTM 950 


OPHIOGLOSSACEAE 

mala ll a atum (Sav.) Underwood; DTM 2012 
preng.; Raymond 1600 NLU 

see Sw. RDT 148118 

Botrychium virginianum (L.) Sw.; DTM 2222 

Ophioglossum crotalophoroides Walt.; MM 6062 


iecocfiy 


Bohn hy / ai. 


LOUISIANA 1201 


ied ee Sng e snail Prantl; RDT 34526 LSU 


udicaule Lf; RDT 5 


1996 
Onhiogiesum petiolatum Hook.; RDT 33845 LSU 
Ophioglossum vulgatul re le MM 1 
OSMUNDACEAE 
Osmunda cinnamomea L., DTM 2156 
Osmunda oe L.; DTM 2157 
POLYPO 
Pleopeltis eee oides (L.) Andrews &Windham;DIM 
949 


SELAGINELLACEAE 

Selaginella arenicola Underwood; MM 2840 
THELY PTERIDACEAE 

Thelypteris kunthii (Desv.) Morton; DTM 1734 
Thelypteris palustris Schott; DTM 1212 


GYMNOSPERMS 


CUPRESSACEAE 
Juniperus virginiana L.; DTM 1609 
odium distichum (L.) L.C. Rich.; MM 6514 
PINACEAE 
Pinus echinata P. Mill; MM 6199 
Pinus taeda L. MM 5982 


ANGIOSPERMS 
MONOCOTYLEDONS 


AGAVACEAE 
Manfreda virginica (L.) Salisb. ex Rose; DTM 1861 
Yucca louisianensis Trel., MM 241 


ALISMATACEAE 

Echinodorus cordifolius (L.) Griseb.; DIM 77 

Sagittaria graminea Michx, DTM 1315 

Sagittaria latifolia Willd.; DTM 672 

*Sagittaria montevidensis Cham.& Schlecht; DTM 1590 
Sagittaria papillosa Buch.; DTM 1300 

Sagittaria platyphylla (Engelm.) J.G.Sm.;DTM 1216 


ARACEAE 

Arisaema dracontium (L.) Schott; DTM 377 
Arisaema triphyllum (L.) Schott; DIM 2153 
*Colocasia esculenta (L.) Schott; DIM 1306 
Peltandra virginica (L.) Schott; DTM 2224 


ARECACEAE 

Sabal minor (Jacq.) Pers.; MM 190 
BROMELIACEAE 

Tillandsia ides (L.) L.; DTM 3002 


BURMANNIACEAE 
Burmannia biflora L.; Reid 4639 LSU (Reid, pers.comm.) 


CANNACEAE 
*Canna indica L., DTM 1855 (questionably naturalized) 


COMMELINACEAE 
Gt lina communis L.; DTM 2329 
Commelina diffusa Burm.f. DTM 1408 
Commelina erecta L.; DTM 141 


ommelina virginica L DIM 486 
Tradescantia hirsutiflora Bush; DTM 
Tradescantia occidentalis (Britt.) fae DIM 473 
Tradescantia ohiensis Raf; DTM 1 ae 
Tradescantia reverchonii Bush; DTM 1 


CYPERACEAE 

Bulbostylis capillaris (L.) Kunth ex C.B. Clarke; RDT 
155761 

Bulbostylis ciliatifolia (Ell.) Fern.; MM 2833 


pe ae icans wal ick ex Spreng.; Raymond 1722 NLU 
chwein.; Hyatt 8580 LSU 

ne arkansana (Bailey) ae ee 25936 LAF 

Carex atlantica ea RDT 95343 LS 

Carex austricina Mackenzie; nov 98507 

Carex planed Dewey; RDT 17006 

Carex 


Schwein; MM ae 


Carex is Schwein.; DTM 1736 
Carex complanata Torr.& Hook.; Barbour 990 
Carex crebriflora Wieg.; Raymond 1234 NLU 
Carex crinita Lam.; DIM 2231 
Carex crus-corvi Shuttlew. ex Kunze; DIM 1353 
Carex debilis Michx.; RDT 95345 
Carex decomposita Muhl.; MM 7429 
Carex festucacea Schkuhr ex Willd.; Thieret 22620 LAF 
Carex flaccosperma Dewey; Raymond 1213 LSUS 
Carex frankii Kunth; DTM 1281 
Carex glaucescens Ell; Raymond 1359 LSU 
Carex glaucoidea be ex Olney; RDT 99544 
Carex hyalina Boott; MM 3 
ae Rae Steud.; ie 1430 NLU 

arex intumescens eee ea 979 
ia Bailey; RDT 
Carex leavenworthii nes i 
Carex leptalea Wahl.; Reid a a (Reid, pers.comm.) 
Carex louisianica Bailey; D 
Carex ee ina Muhl. ex Re an D352 
Ca arex lurida W Nahlenb.; DTM 1 236 
Carex meadii Dewey; MM 2525 
Carex muehlenbergii schkuhr ex Willd.; RDT 88719 
Haynes 5697 


Se anne Naczi; Raymond 1677 NLU 
s (Bailey) Small; ae 1214NLU 
Carex retrolexa DE ex ee rad 


LI 
UICA style HCAU DULKI 


Carex texensis (Torr.) Bailey; i 88223 

Carex triangularis Boeckl.; RDT 99524 

Carex tribuloides Wahlenb.; : 70062 

Carex vulpinoidea Michx.; D 349 

Cyperus acuminatus Torr. & Hess ex Torr.; DTM 2332 
Cyperus compressus L.; RDT 32983 

Cyperus croceus Vahl; MM 6834 


1202 


*Cyperus difformis L.; MM 6993 

Cyperus echinatus (L.) Wood.; DTM 1931 

Cyperus erythrorhizos role oo 5430 

Cyperus esculentus L.; MM 6857 

Cyperus filiculmis Vahl; 170174 (Ferguson, pers. 
comm.) 

ae os L.; CE. DePoe 6652-1 

m.) 
eo ee ee Shireman 57 LSU 
ici iria L.; DTM 
Cyperus odoratus L.; nee 

Cyperus oxylepis Nees ex on RDT 37207 

Cyperus plukenetii Fern.;Lynch 2639 LSUS 

Cyperus polystachyos Rottb.; Lynch 2674 LSUS 

Cyperus. pseudovegetus Steud.; RDT 166992 (Ferguson, 
pers.comm.) 


NLU (Fergu- 


Cyperus retroflexus Buckl.; MM 2725 

Cyperus retrorsus Chapm.; MM 6630 

*Cyperus rotundus L.; DTM 1361 

Cyperus strigosus L; RDT 45424 (Ferguson, pers.comm.) 
Cyperus virens Michx.; Overby 85 NLU (Ferguson, pers. 


comm.) 
Eleocharis acicularis (L.) Roemer & J.A. Schultes; DTM 
1458 
Fleocharis obtusa (Willd.) J.A. Schultes; DTM 1238 
Eleocharis palustris (L.) Roemer & J.A.Schultes; MM 6429 
Eleocharis parvula (Roemer & J.A. Schultes) Link; Bar- 
bour 498 


Eleocharis tortilis (Link) J.A.Schultes; RDT 33624 
Eleocharis tuberculosa (Michx.) Roemer & J.A.Schultes; 
T 36268 


Fimbristylis autumnalis (L..) Roemer & J.A.Schultes; DTM 
1567 


Fimbristylis miliacea (L.) Vahl: DTM 1542 
Fimbristylis puberula (Michx.) Vahl; ete 2147LSUS 
Fimbristylis tomentosa Vahl; DTM 
Fimbristylis vahlii (Lam.) Link; a 2 
Fuirena simplex Vahl: MM 6673 
me valle Michx.; RDT 14068 
Hook. & Arn. ex Torr,; ov 2829 
Isolepis molesta (M.C. Johnston) S.G. Sm.; Shinners 
28087 BRI 


Kyllinga cae Rottb., DIM 1928 
yllinga odorata Vahl; DTM ae 

Kyllinga pumila Michx.; RDT 3 

Lipo ae micrantha (Vahl) [ ae Shireman 56 


~_ 


ep caduca Ell; Raymond 2028 LSUS 
hospora corniculata (Lam.) A.Gray; DTM 1113 

dee sic glomerata (L.) Vahl; Barbour 494 

Rhynchospora harveyi W. Boott; MM 7427 

Rhynchospora inexpansa (Michx.) Vahl; Haynes 5352 

Scirpus atrovirens Willd.; Allen 7975 NO 

Scirpus cyperinus (L oe 1448 

Scleri eG ata MCN: RDT 

Scleria tt ichx.; ne — LAF 


BRIT.ORG/SIDA 22(2) 


Sc leria paucinora Muhl. ex Willd.; Allen 9879 LAF 


Creel ChIQGUIOT TICE Michx.; RDT 92403 
Websteria confervoides (Poir.) S. Hooper; Barbour 495 


DIOSCOREACEAE 

Dioscorea villosa L; DTM 1288 

HYDROCHARITACEAE 

*Egel ia densa Plat ich.; Barbour 483 

Limnobium spongia (Bosc) L.C. Rich. ex Steud.; Haynes 
5342 


IRIDACEA 
Alophia ae mondii (Graham) R.C. fe oo 1762 
tia lahue (Molina) Goldblatt; M 
ris brevicaulis Raf; Raymond 2094 ne 
: fulva Ker-Gawl.; French s.n. NLU 
Iris giganticaerulea Small; Overby 192 NLU 
“ris pallida Lam.; Laborde 44 
*Iris pseudacorus L.; MM 172 
Iris virginica L.. DTM 2291 
Nemastylis geminiflora Nutt; DTM 369 
eats aaa ifolium P.Mill; DTM 415 
ickn.; DIM 340 
Sisyrin ich jum langloissii aie MM 2558 
Sisyrinchium minus Engelm.& A. ee a 25939 LAF 
Sisyrinchium rosulatum Bickn.; DTM 


JUNCACEAE 

Juncus acuminatus Michx.; RDT 81369 

Juncus brachycarpus rae ;DIM 1112 

Juncus bufonius L.;M 

oo capitatus ne oan 128670 LSU 
INCU M nzie; DePoe 6650 NLU 

Juncus dichotomus Ells D DTM 1272 

Juncus diffusissimus Buckl.; DTM 1234 

J 


Juncus marginatus Rostk.; DTM 1235 
Juncus nodatus Coville; Raymond 1241 NLU 
Juncus polycephalus Michx.; RDT 629 
Juncus repens Michx.; RDT 136859 
Juncus scirpoides Lam.; RDT 33633 

Juncus tenuis Willd.; DTM 1163 
Juncus torreyi Coville RDT 128675 

uncus validus Coville; DTM 126 

Luzula bulbosa (Wood) Smyth & Smyth; Haynes 4568 
LEMNACEAE 

Lemna aequinoctialis ew ; Overby 86 NLU 
Lemna minor L.; Hayne 


Lemna valdiviana Phil. ae 


Spirodela polyrrhiza (L.) Schleid.; Haynes 4155 
Spirodela punctata (G.F.W.Mey.) C.H. Thompson; Haynes 
422 


Wolffia brasiliensis Weddell; Haynes 41 f 
Wolffia columbiana Karst.; Overby 227 
Wolffiella gladiata (Hegelm.) Hegelm.; me 4153 


LOUISIANA 1203 


LILIACEAE 


Alliitm canandense| « 


TM 3058 
ce nena L.; MM s.n (questionably natural- 
d). 


ze 
Erie scilloides (Raf) Cory; DTM 3015 

Cooperia drummondii Herbert; a oi 

Erythronium ies Nutt; DTM 2 

Habranthus tubispathus (L'Her.) ae MM 262 
*Hemerocallis fulva (L.) L.; DTM 1723 (localized and 


persisting) 
Hymenocallis caroliniana (L.) Herbert; DTM 1843 
Hymenocallis liriosome (Raf.) Shinners; DTM 853 
Hypoxis hirsuta (L.) Coville; DTM 343 (includes H. rigida 


Chapman) 
*Narcissus pseudonarcissus L.; Barbour 559 (localized 
and persisting) 
Nothoscordum bivalve (L.) Britt; DTM 250 
*Nothoscordum inodorum (Aiton) Nichols; DTM 1675 
*Ornithogalum umbellatum L; DTM 3014 
Polygonatum biflorum (Walt.) Ell; DTM 674 
Schoenolirion wrightii Sherman; MM 7291 
Stenanthium gramineum (Ker-Gawl.) Morong; DTM 
811 
Trillium recurvatum Beck; MM 144 
Trillium pusillum Michx,; DTM 2185 
Uvularia sessilifolia L.; DTM 2617 
*Zephryanthes candida (Lindl.) Herbert; Barbour 467 
nee nuttallii (Gray) S.Wats.; DIM 861 


NAJADACEAE 
Najas guadalupensis (Spreng.) Magnus; DTM 1522 


ORCHIDACEAE 

Corallorhiza wisteriana Conrad; DIM 11 

pagcnata Tepes Nutt; ar 1947 i 

rid © ied og Ge cillata Raf. -DTM 

ea australis Lindl; DTM a 

Malaxis unifolia Michx.; RDT 43537 

Platanthera ciliaris (L.) Lindl; RDT 42439 

Platanthera clavellata (Micha Luer; MM 256 

flava (L.) Lindl; Raymond 1483 NLU 

Platanthera lacera ae G.Don; DTM 1759 

Spiranthes cernuus (L.) L.C.Rich.; MM 278 

Spiranthes lacera (Raf.) Raf; Overby 262 NLU 

Spiranthes laciniata (Small) Ames; DTM 2013 

Spiranthes odorata oe ee ie 5426 

Spiranthes ovalis Lindl; 

Spiranthes praecox bevy - vo DTM 2313 LSU 

Spiranthes tuberosa Raf.; DT 

Spiranthes vernalis Engelm. & iy DIM 903 

ae aria goer) NO aNens oe LSU 
R ; 1529NLU 


P latanthera 


=> 


POACEAE 

Agrostis elliottiana J.A. Schultes; DTM 1632 
*Agrostis gigantea Roth; DTM 12 

Agrostis hyemalis (Walt.) B.S.P.; DTM 2029 
Agrostis perennans (Walt.) Tuckerman; DTM 1416 


Agrostis stolonifera L DTM 1309 

*Aira caryophyllea L., DTM 3059 

*Aira elegans Willd.ex Kunth; DTM 1659 

Alopecurus carolinianus Walt.; DIM 2603 

Andropogon gerardiiVitman; DTM 1458 

Andropogon glomeratus (Walt.) B.S. i DTM 1498 

A OPER ON ternarius Michx.,, DTM 

Andropogo ia virginicus L.; ‘DTM 1 ys 

el KCN, qustatae Boiss.; DIM a 
Trin. & Rupr; MM 28 


Aristida dichotoma Michx.; DTM 2004 

Aristida lanosa Muhl. ex Ell; MM 2928 

Aristida ene eee fies DTM 1563 

Aristida MM 2893 

Aristida purpurascens ie DTM 1468 
Arundinaria gigantea (Walt.) Muhl.; DTM 2193 
*Arundo don 1569 


*Avena fatua L.; DTM 1732 

*Avena sativa. DIM 2615 

Axonopus fissifolius (Raddi) Kuhlm.; DTM 1264 

*Bothriochloa ischaemum (L.) Keng; RDT 68561 

Bothriochloa languroides (DC.) Herter; DTM 1382 

Bothriochloa longipaniculata (Gould) Allred & Gould; 
DTM 2079 


se springfieldii (Gould) Parodi; Thieret 24481 


res iza minor. DTM 
*Bromus catharticus ae DTM 2298 
*Bromus diandrus Roth; RDT 65106 LSU 


Buchloe dactyloides (Nutt.) coer Snyder s.n.US 
Cenchrus spinifex Cav.; MM 2 
Chasmanthium latifolium ies DTM 1735 
Chasmanthium laxum (L.) Yates; DTM 1384 
Chasmanthium ee florum (Poir.) Yates; DTM 2557 
Chloris virgata Sw.;D 320 
*Cynodon dact ctylon (L NY Pers.; one 4237 


ad 
< 
aa 
iS) 
n 
cy 
= 
cS 
a 
tas) 
in 
aed 
= 
g 
mi 
ey 
Wn 
is 


*Dactylis glomerata L.,; DTM 1 
*Dactyloctenum aegyptium He ine Overby 121 NLU 
Danthonia spicata (L.) Beauv.ex Roemer &J.A.Schultes; 


M 1655 
Dichanthelium aciculare (Desv. ex Poir.) Gould & CA. 
Clark; DTM 1671 
Dichanthelium acuminatum (Sw.) Gould & C.A. Clark; 
DTM 1210. 
Dichanthelium boscii (Poir.) Gould and C.A. Clark; DTM 


1408 
Dichanthelium commutatum (J.A. Schultes) Gould; 
DTM 1241 


1204 
Dichanthelium depauperatum (Muhl.) Gould; Lewis 
NLU 
Dichoanthelittm dic-hantamirim (I L.) Gould; DTM 1 182 
m.) Gould; DTM 1239 

sec sca linearifolium ee ex Nash) Gould; 
Allen 

Pe Sosa oligosanthes (J.A.Schultes) Gould; MM 
2617 

Dichanthelium ovale (Ell.) Gould & C.A.Clark; Lewis 3585 
N 


Nichanthel lavifl 


nthes Schult.; MM 6491 
a a ieveneln (Scribn. & Merr.) Gould; DTM 


ichanthelitim polya 


Dhan boasted item) Gould; DTM 1403 


Dich Ell) Gould; DTM 1274 
cane on (Retz) Koel: D M1414 
Digitaria cognata (JA. aaa Pilger; DTM 1527 


Digitaria filiformis (L.) Koel, DTM 1544 
*Digitaria ischaemum (Schreb.) Schreb. ex Muhl; DTM 


*Digitaria violascens Link; DTM 137 a 
*Echinochloa colona (L.) Link; DTM 1 
*Echinochloa crus-galli (L.) Beauv.; oe 277 SU 
Echino chloa muricata (Beauv.) Fern.; MM 6854 
Echinochloa walteri i (P urs| 1) Heller; DTM 2553 
*Eleusine indica (L (L.) Gaertn.; DTM 1596 

Elymus nicus L.; DTM 2999 
pri ees (All.) Vign. ex Janchen.; DTM 1420 
*Eragrostis curvula (Schrad.) Nees; DTM 2312 
Eragrostis hirsuta (Michx.) Nees; MM 2927 
Eragrostis hypnoides (Lam.) B.S.P; DTM 1479 

*Eragrostis japonica ea Trin; DTM 2706 
Eragrostis lugens Nees; DTM 1643 

*Eragrostis minor Host; ae 1377 

*Eragrostis pilosa (L.) Beauv.; DTM 1510 
Eragrostis refracta (Muhl.) Scribn.; DTM 1997 
Eragrostis reptans (Michx.) Nees; DTM 1986 
Eragrostis secundiflora J. Presl; DTM 1803 
Eragrostis spectabilis (Pursh) Steud.; MM 2917 
man trichodes (Nutt.) Wood; MM 2918 

hloa ophiuroides (Monro) Hack,; DTM 1363 

ie acuminata (J. Pres!) Kunth; DTM 1971 
Eriochloa contracta A.S. Hitchc.; DTM 1481 
Festuca paradoxa Desv., DTM 1277 

te cisanana Fern. OM 2327 
ichx.) B.S.P; DTM 1489 


*Holcus lanatus L.; DIM 1346 

Hordeum pusillum Nutt. DTM 1621 

Leersia lenticularis Michx.; Haynes 5422 

Leersia oryzoides (L.) Sw., DTM 1488 

Leersia virginica Willd; DTM 1359 

Leptochloa fusca (L.) Kunth; DTM 1325 

Leptochloa panicea (Retz.) Ohwi; MM 6767 

Leptochloa panicoides (J. Presl) A.S. Hitche. DIM 1 

*Lolium arundinaceum (Schreb.) S.J. Derbyshire; 7 
139] 


BRIT.ORG/SIDA 22(2) 


*Lolium perenne L.; DIM 1187 
*Lolium temulentum L; DTM 1673 
Luziola fli itans (Michx.) Terrell & Robins., DTM 1549 
Melica mutica Walt.; DTM 1614 
Muhlenbergia schreberi J.F.Gmel.; 
Nassella leucotricha (Trin. & Rupr.) an oa 3050 
Oplismenus hirtellus (L.) Beauv.; ca 1305 
Panicum anceps Michx,; DTM 1 
Panicum brachyanthum net au 1998 
Panicum capillare L.; DTM 
Panicum dichotomiflorum rie DIM 1386 
Panicum flexile (Gattinger) Scribn.; Correll & Correll 

10097 LSU 
Panicum eet Ell.; Haynes 5425 
Panicum A.Schultes; RDT 140681 
He sc.ex Nees; DIM 139 
Panicum verrucosum Muhl.; ae 1529 
Panicum uoene L; DTM 
atatum Poir.; a 
t m (L) L, rae fee 
Paspal listich L.; DTM 1970 
Paspalum floridanum Michx.; DTM 1389 
uncial ave lleNs DIM 1374 

i (Fourn.) Nash; DTM ny 

Fluegg ge; DIM 1 


Paspalum pli atriltim NAj hx.; DTM oo 


ss 


Pan j m rigid 


es ror 


palum TEPIOCOCUT YI 


Paspalum pubi florum Rupr. ex Fourn.; DTM 1586 
Paspalum repen Bera. R 47 
ean ara Bn . igs 

villei Steud.; DTM 1 
phalaris gare iniana ae DIM ee 
*Phleum piaineh L.; RDT 65054 
*Phyllosta a Carr ex A. Riv.& C. Riv, MM 7244 
Pipt (L.) Parodi; DTM 2297 
*Poa annua L.; DTM 1 
Poa autumnalis the ex Ell; DTM 2186 
Poa pratensis L.; Thieret 22666 LAF 
*Polypogon monspeliensis (L.) Desf; RDT 66455 LSU 
mean pe ranse ald (L.) Nutt. at 2959 


haetitim aqvengq 


winii Spreng.;RDT 3727] 

ee. by evi arbe (Michx.) Pers. DTM 1469 
h teum (Walt.) Pers.; DIM 1536 

Sacci ek sia (L.) Nash; DTM 1431 
Schedonnardus paniculatus (Nutt.) Trel.; DTM 2627 
ee aoe scoparium (Michx.) Nash; DTM 1533 
*Secale cereale L., DIM 2232 
*Setaria faberi Herrm.; RDT 119026 
Setaria parviflora (Poir.) Kerguelen; MM 2055 
*Setaria pumila (Poir.) Roemer &J.A.Schultes; DTM 1573 
*Setaria viridis (L.) Beauv.; RDT 45434 
Sorghastrum elliottii (Mohr) Nash; MM 2931 
Sorghastrum nutans (L.) Nash; DTM 2005 
*Sorghum bicolor (L.) Moench; DTM 1411 
*Sorghum halepense (L.) Pers.; Haynes 4228 
Sphenopholis intermedia (Rydb.) Rydb.; RDT 88229 
Sphenopholis nitida (Biehler) Scribn.; DTM 1626 


— 


MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH, LOUISIANA 1205 


Sphenopholis obtusata (Michx.) Scribn.; DTM 2632 
Sporobolus clandestinus (Biehler) A.S. Hitche.; RDT 
37044 


Sporobolus compositus (Poir) Merr; DTM 1579 
pee eee tus peana (Torr.) A.Gray; DIM 1578 
I ;DT 
! 


coerce us junceus Gesu) Kunth; DTM 1515 

Sporobolus pyramidatus (Lam.) Hitche.; MM 7277 

Sporobolus vaginiflorus (Torr.ex Gray) Wood; RDT 37396 
LSU 


Steinchisma hians (Ell.) Nash; DTM 1 

Stenotaphrum secundatum (Walt.) nc rs 308 
Tridens ee (Ell.) A. Schultes; DTM 

Tridens flavus (L) A.S.Hitche.,MM 2915 oa Gee 


vai Hy 
chapmanii (Small) Chase 


136 LSUS 
ve 2932 


Trisetum interruptum Buckl.; Allen 3774 NLU 

*Triticum aestivum L.; DTM 2645 

Urochloa platyphylla (Monro ex Wright) R.Webster; 
DTM 1378 


*Urochloa ramosa (L.) Nguyen; DTM 13 
Urochloa texana (Buckl.) R Webster; iene Burkett 


Co. s.n. LSU 

*Vulpia bromoides (L.) S.F.Gray; DTM 1228 

ve a Leyines (L.) K.C.Gmel.; DTM 1650 

(Walt.) Rydb,; DTM 2226 

Vulpia a sciurea (Nutt). Henr.; MM 2537 

Zizaniopsis miliacea (Michx.) Doell. & Aschers.; DTM 
1158 


PONTEDERIACEAE 
*£ichhornia crassipes (Mart.) Solms; DTM 240 
Pontederi data .; DIM 88 


POTAMOGETONACEA 

poor Nee ce Barbour 519 

Potamoge 5L;DIM 1521 

SMILACACEAE 

Smilax bona-nox L.; ale eae 

Smilax glauca Walt.; D 

Smilax laurifolia L: a 25 LSUS 
/ 


smi ax rotund ifolia L.; ab 959 
Smila eri Mii M T 155720 
Smilax tamnoi des 2 ee 32815 
SPARAGANIACEAE 

Sparganium americanum Nutt.; DTM 1830 


gee 


is Pers; DTM 1758 
Wee auiolia L.; DTM 1299 
XYRIDACEAE 

Xyris caroliniana Walt.; Barbour 1099 
Xyris jupicai L.C. Rich.; MM 263 


DICOTYLEDONS 


ACANTHACEAE 
Dicliptera brachiata (Pursh) Spreng.; DTM 227 
Justicia ovata (Walt.) Lindau; DT De 

*Ruellia brittoniana Leonard; RD 

Ruellia caroliniensis (J.F. ee ) ee a 64 
Ruellia humilis Nutt; DTM 8 

Ruellia pedunculata Torr. ex on DTM 447 
ACERACEAE 

Acer barbatum Michx.; RDT 166956 

Acer negundo L.; Haynes 4146 

Acer rubrum L.; Haynes 4583 
Acer saceniaenua ee Sais 2 LSU 
Ac 


AIZOACEAE 
esuvium it (Walt.) B.S.P; Thieret 26136 LAF 
Trianthema portulacastrum L.; DTM 174 
AMARANTH 
“Aenanthea ice ies oo . Griseb.; DIM 84 
Amaranthu i RDT 6 
Amaranthus arenicola |.M. lens RDT 59959 


Amaranthus aie idus LD 
Amaranthus palmeriS. ie oa 1964 


pe spinosus L.; 924 

s viridis La MM 6978 
pa foeeee es ae DTM 1793 
Froelichia gracilis (Hook.) Moq.; RDT 76696 


ANACARDIACEAE 

Rhus aromatica Aiton; DTM 2181 

Rhus copallinum L; DTM 1924 

Rhus glabra L; RDT 33565 

Toxicodendron radicans (L) Kuntze; Haynes 4140 

ANNONACEAE 

Asimina parviflora (Michx.) Dunal; RDT 80926 

Asimina triloba (L.) Dunal; Hardy 8685 

APIACEAE 

Ammoselinum butleri (Enelm.ex S.Wats.) Coult.& Rose; 
RDT 88289 


aaa incana Ruiz & Pavon; ca 7180 
otundifolium L.; RDT res) 
ee tainturieri Hook, DTM 
Se cea (Pers.) ee ex Britt.& 
Wilson; Haynes 577 
Cicuta maculata L,; eee ae ee 
*Conium maculatum L.; DTM 
SECU: eauaGense ve 2 “ 25312 
cyn 


DTM 1190 


*Daucus carota L. DIM an 

Daucus pusillus Michx.; MM 2653 

Eryngium hookeri Walp.; DTM 143 

Eryngium prostratum Nutt.ex DC; Haynes 4109 


1206 


Eryngium yuccifolium Michx.; DTM 142 
*Falcaria vulgaris Bernh.; RDT 88299 
aecayc ranunculoides Lf; sai 1097 


yan Rous umbellata L.; “DIM 


Hydrocotyle verticillata Thu nb.; a 75 


Limnoscadium pinnatum (DC.) Mathias & Constance; 


DIM 478 
Oxypolis rigidior (L.) Raf.; ROT 7 
Polytaenia nuttallii DC.; DTM 549 
Prilimmnium capillaceum (Michx.) Raf.; DTM 1022 
(Ell.) Raf; RDT 128660 
im nuttallii (DC.) Britt.; Raymond 770 LSUS 
Sanicula canadensis L.; DTM 1040 


— 


Dt ilire 


Prilingniy 
I 


Sanicula odorata (Raf.) K.M. Pryer & L.R. Philliope; RDT 
33075 


Sanicula smallii Bickn.; RDT 170095 
Spermolepis divaricata (Walt.) Raf.ex Ser; DTM 2285 
et a echinata (Nutt. ex DC) Heller; Allen 7999 


Siar een (Nutt.ex DC.) Mathias & Constance; 


Taenidia integerrima (L.) Drude; Cocks s.n. NO 
*Torilis a is (Huds.) Link; DTM 29 

*Torilis noueya ey DTM 1033 

*Tre) rpu je Nutt.ex DC, DTM 464 
Zizia aurea (L.) W.D i Koch; Raymond 732 LSUS 


APOCYNACEAE 
eee nae he he 838 
LMM 


recs difforme i: ) " a DIM 454 
*VINC 

ne minor Ls RDT ns 

AQUIFOLIACEAE 
llex ambigua (Michx.) Torr; RDT 140651 LSU 
llex decidua Walt.; Haynes 4093 

Ilex opaca Aiton; Hardy 8616 

Ilex vomitoria Aiton; DTM 1151 
ARALIACEAE 

Aralia spinosa L.; DTM 1154 
ARISTOLOCHIACEAE 

Aristolochia reticulata Jacq.; RDT cue 
Aristolochia serpentaria L; RDT 119103 
Aristolochia tomentosa Sims; RDT 36969 
ASCLEPIADACEAE 

Asclepias amplexicaulis Sm.; DTM 2294 
Asclepias perennis Walt., DTM 523 

fog ne tuberosal.; DTM a 

riegata Le -DIM8 


Asclepias Vu 
Asclepias verticillata L.; DTM ve 
siete viridifiora Raf; DTM 556 


idis Walt.; DIM 62 


Asclepias Vil 
Cynanchum laeve (Michx.) Pers.; Hardy 6382 
Matelea cynanchoides (Engelm.) Woods.; MM 2613 


BRIT.ORG/SIDA 22(2 


Matelea decipiens (Alexander) Woods.; RDT 51975 
Matelea gonocarpos (Walt.) Shinners; RDT 36994 


ASTERACEA 
ACNE mill flu i. seal 329 
m.) R.K. Jansen; DTM 442 
Ageratina ‘alti issima (L ) ce & E.Robins.; DIM 693 
Ambrosia artemisiifolia L., DTM 9 
Ambrosia psilostachya re si in 
Ambrosia trifida L DTM 
nears i Hayes 
nthemis cotula L.; D 
nen. cali a )Trel.; DTM 1771 
be eal po neum Raf., DTM 407 


lia L.; Hardy 6871 


” 


Berlandiera betonicifolia (Hook.) Small; DTM 1074 
Berlandiera pumila (Michx.) Nutt. DTM 997 
Bidens aristosa (Michx.) Britt., DTM 1993 

Bidens bipinnata L.; Haynes 5419 

Bidens discoidea (Torr. & ie Britt; DTM 1 
Bidens frondosa L.; DIM 
Bidens laevis (L.) B.S.P; ae os 
Boltonia di Ell; DIM 718 
*Calyptocarpus vialis Less.; DTM 1860 
*Carduus nutans L.; DTM 460 
Chrysopsis pilosa — oie 2720 
#Cj horinm intyl ni 223 
Cirsium alti es DIM 5 
Cirsium eee (Walt.) Fern. & oe DTM 840 
Cirsium engelimanill ae cen LSU 

,DIM5 


Cirsium 


Conoclinium coelesti (L.) DCD 639 

*Conyza bonariensis (L.) Cronq.; a 321 

Conyza canadensis (L.) Cronq.; DTM 589 

Coreopsis grandiflora Hogg ex Sie DIM 434 

Coreopsis intermedia Sherff; MM 2605 

Coreopsis lanceolata L.; DTM 2 a 

Coreopsis tinctoria Nutt.; Raymond 768 LSUS 

*Crepis pulchra L.; DIM 842 

Croptilon divaricatum (Nutt.) Raf.; DTM 592 

Doellingeria sericocarpoides Small; DTM 1947 
copis amplexicaulis (Vahl) Cass.; DTM 60 


59 LAF 


) 


Dra 
Echinacea pallida (Nutt.) Nutt; DTM 491 (includes £. 


angustifolia DC. and E. sanguinea Nutt.) 
Echinacea purpurea (L.) Moench; RDT 136864 
Eclipta prostrata (L.) L., DTM 247 
Elephantopus carolinianus Raeusch.; DTM 8 
Elephantopus nudatus Gray; RDT 155792 

diate oe es tosus L.; DTM 241 
an Raf. ex DC; DTM 1524 
DTM 1988 

oe ae ie DIM 45 
Erigeron pulchellus Michx.; Raymond 750 LSUS 
Erigeron strigosus Muhl.ex Willd, DTM 52 
oe tenuis ae & A.Gray; DTM 360 
(Lam.) Small; DTM 2007 


Eriger Nannie (l .) Pe 


LOUISIANA 1207 


Eupatorium compositifolium ea ;DTM 2052 
Seana ad L;DTM 
indifolium L a 


ynoatoris 1m 


Eupatorium semiserratum DC.; ay ah (includes E. 
glaucecsens Ell. 

Eupatorium serotinum Michx,; DTM 172 

Eurybia hemispherica (Alexander) Nesom; DTM 705 

=e leptocephala (Torr. & A. Gray) Greene; DTM 


7 


i een & A. Gray) A. Gray; al a 1 LSU 
*Facelis retusa (Lam.) Schultz-Bip.; DIM 8 
Fleischmannia incarnata (Walt.) King & ie aie RDT 


33060 
Gaillardia aestivalis (Walt.) H. ines DTM 1800 
Gaillardia pulchella Foug.,, DTM 
Gamochaeta te (Urb.) eee DTM 400 
Gamochaeta a a Nesom; Strout 112 LSUS 
Gamochaeta pers (Willd.) Cabrera; DTM 1751 
Gamochaeta purpurea (L.) Cabrera; DTM 373 
Helenium amarum (Raf.) H. Rock; DTM 24 
Helenium flexuosum Raf.; DTM 27 
BISNanENtS alc L;DIM 178 
arbo a 


LE 


Helianthus eee L; DTM 1894 
pea ae ee DTM 1756 
a 


elianthus mollis Lam.; -RDT 362 
lianthus occidentalis Riddell; oe. s.n.NO 
Helianthus petiolaris Nutt.; Se s.n.NO 
Helianthus strumosus 
Heterotheca subauians oe ) a & Rusby; MM 2777 
Hieracium gronoviiL.; DTM 390 
Hymenopappas a He DC.; Barbour 1193 
Hymenopappas scabiosaeus L'Her,; DTM 17 
*Hypochaeris glabra L.; DIM 2830 
lva angustifolia Nutt. ex DC; DTM 2077 
lva annua L.; DTM 2025 
Krigia caespitosa (Raf.) Chambers; DTM 38 
Krigia dandelion (L.) Nutt; Raymond 731 LSUS 
Krigia occidentalis Nutt.; DT 
Krigia virginica (L.) Willd; ae - 3 
Lactuca eqnadens L;DTM 1 
Lactuca floridana (L.) Gaertn.; ar 651 
Lactuca Seine oe hia RDT 36155 
*/ actuca serriola L.; DIM 4 
*Leucanthemum ae DTM 1 
naturalized) 

Liatris aspera Michx.; DTM 
Liatris elegans (Walt.) Ante ane 204 
Liatris pycnostachya Michx.; DTM 517 
Liatris squarrosa (L.) Michx,, DTM 542 
Hale sala is shinners; RDT 37304 
ex DC.; DTM 351 
Mba a scandens ( . 2 ae DTM 209 

li rr. & A. Gray) Small; DTM 1 
Olig igoneuron rigidum (| ic Small; RDT 93593 


ye 
e 


19 (questionably 


Packera gla bella (Poir. ) G, Jeffrey; DIM 41 
Packera ahora (Muhl.ex Willd.) W.A.Weber & A. Love; 
2 


*Parthenium hysterophorus L.; DTM 1999 
Parthenium intearifolium Ee *-DIM 13 
Pityopsis ee (Michx.) Nutt.; MM 906 
eacampho cE ;DIM 698 
Pluchea fecal soe MM 284 

Pluchea odorata (L.) Cass.; MM 277 
Pseudognaphalium helleri (Britt.) A.Anderb.; RDT 41805 
Pseudognaphalium obtusifolium (L.) Hilliard & Burtt; 


20738 
= 
a 


Pyrrhopappus carolinianus (Walt.) DC.; DTM 552 

Pyrrhopappus pauciflorus (D. Don) DC; DTM 499 
atibida pinnata (Vent.) Barnh.; DTM 94 

Rudbeckia grandiflora (D.Don) J.F. Gmel. ex DC.; DTM 


Rudbeckia hirta L; DTM 1 
Rudbeckia maxima Nutt,; ow 95 
*eneck vulgaris L.; MM 5 
aad um He ifolium Niche DTM 2326 
inia 20 


Inhitwym tum L.; 


*Sily bum marianum (L ) ce RDT 345 


small anthus uvedalia (L.) Mackenzie ex ad DTM 


1088 
Solidago altissima L.; DIM 627 
Solidago auriculata Shuttlew. ex Blake; DTM 1916 
aie udoviciana \Gley) ee MM 2889 
odora Aiton 
Soop eto ane Aiton: oe 96 1 
Solid nes 4081 
20 bse cages B Mil iime 686 
hl ex Willd.; DTM 670 
Soli iva eos Ruiz & Pavon; DTM 1014 
*Sonchus asper (L.) Hill; DIM 424 
*Sonchus oleraceus L.; DTM 896 
si aes trichum divaricatum (Nutt.) Nesom; Haynes 
4261 


> 
a 


— richum drummondii (Lindl.) Nesom; DTM 


ee a trichum dumosum (L.) Nesom; MM 6957 

Symphyotrichum laeve (L.)A.& D. Love; DTM 744 

Symphyotrichum lanceolatum (Willd.) Nesom; DTM 
2586 


Symphyotrichum lateriflorum (L) A.& D.Love; DTM 2057 
Sp euenum cee (Wieg. ) Neon RDT 69310 
Riddell DIM 


' 


b 


NANA BO00 


570 
Symphyotrichum patens (Aiton) N 
Symphyotrichum pilosum (Willd.) econ DTM 2569 
Symphyotrichum praealtum (Poir.) Nesom; DTM 700 
Symphyotrichum pratense (Raf.) Nesom; DTM 2080 
Symphyotrichum racemosum (Ell. 
Symphyotrichum subulatum (Michx.) — Hardy 

5297 


NI -MM QOQ0Q 


a 


*Taraxacum officinale G.H.Weber ex Wiggers; DTM 283 


1208 


pa ero Cone (Torr. & A. Gray) A.Gray ex 
Hall; M 
Plea 4 a lium (Hook.) A. Gray; ony 88997 LSU 
Verbesina helianthoides Michx.; DTM 4 


Verbesi Hi virginica L; ‘ Barbo sts 
Vernonia baldwinii Torr; MM 6 
Vernonia missurica Raf; amet 773 LSUS 


Vernonia texana (Gray) Small; Ane 126 


anthium strumarium L., DTM 1095 
ie ica (L.) DC; DTM 3012 
BALSAM 


ACEAE 
ae alee Meerb.; DTM 1094 


BERBERIDACEAE 

*Nandina domestica Thunb; MM 6465 (questionably 
naturalized) 

Podophyllum peltatum L., DTM 47 


BETULACEAE 

Alnus serrulata (Aiton) Willd.; DTM 2228 

Betula nigra L.; Hard 

Carpinus carolin iana Walt; Haynes 5316 

Ostrya virginiana (P. Mill) K. Koch; Haynes 4569 

BIGNONIACEAE 

Bignonia capreolata L.; DIM 803 

Campsis radicans (L.) Seem. ex Bureau; DIM 137 

Catalpa bignonioides Walt.; Barbour 82 

Catalpa speciosa (Warder) Warder ex Engelm.; Frank 
134 LSUS 


BORAGINACEAE 
*Bugl Tal nsis s (L.) .M.Johnston; DTM 1071 
baa 132 LAF 


7 
Holint 


He fotopium indicum i DIMI 
He jotropium pro rmbens ae a 624 


Lith nerm! \A/a| 


citmospermum Cal nse (Walt. ex JF. Gmel.) M acM.:; 


MM 2534 
Myosotis macrosperma Engelm.; MM 6155 (includes 
osotis verna Nutt.) 
Onosmodium bejariense DC. ex A.DC.; Cocks s.n. NO 


BRASSICACEAE 

*Arabidopsis thaliana (L.) Heynh.; RDT 70506 
*Camelina microcarpa DC.; RDT 65101 LSU 

*Capsella bursa-pastoris (L.) Medik.; DTM 256 
Cardamine bulbosa (Schreb. ex Muhl.) B.S.P; MM 6205 
x.) Sw; MM 143 


Cardamine COMCALEN GLA (Michx 

*Cardamine hirsuta L; DTM 261 
Cardamine parviflora L.; Barbour 1133 

Cardamine pensylvanica Muhl.ex Willd; DTM 1 
*Coronopus didymus (L.) Sm. DIM 852 

*Descurainia sophia (L.) Webb. ex Prantl; RDT 65110 

ie brachycarpa Nutt. ex Torr.& A. Gray; DTM 2587 

ifolia Nutt. ex Torr. & A. Gray; DTM 943 
Lepidium den n Schrad.; RDT 65049 LSU 
Lepidium VIFGINIC um Ls ;Raymond oe LSUS 


*Rapistru mM rliqos mit L.) All; DIM 


¥R ; cr ry 
PP ACU 


ie Hayek; DTM 2204 


qatnatrictym 
AM-aqguauiculn 


BRIT.ORG/SIDA 22(2) 


Rorippa palustris (L.) Bess.; DTM 975 
Rorippa sessiliflora (Nutt.) AS. Hitch.; Barbour 970 
36 


Sibara virginica (L eens 
as arvensis iL, DTM 27 

L,; ieee 
*S) ffici (L.) Scop; RDT 83089 


oe ae ae Hook.; MM 2656 
*Thlaspi arvense L.; Barbour 1003 


BUDDLEJACEAE 
Polynremitm nr / 


= 


mbens L.;DTM 154 
CABOMBACEAE 

Cabomba caroliniana Gray; DTM 1721 
CACTACEAE 
Opuntia humitu 


a (Ra f.) R 


Raf; MM 239 


CALLITRICHACEAE 
Callitriche Metron yi Pursh; si 4577 


Callitriche COhPOOU 1S Raf. 4 


CAMPANULACEAE 
Lobelia appendiculata A. ae : TM 732 
Lobelia cardinalis L.; D 
Lobelia puberula Michx.; ae a 5 
Triodanis biflora (Ruiz & Pavon) rae — 387 
[riodanis perfoliata (L.) Nieuwl,; DTM 9 


CAPPARACEAE 
leome hassleriana Chod,; Haynes 5938 
Polen ia dodecandra (L.) DC; RDT 64990 


APRIFOLIACEAE 
*Lonicera japonica Thunb.; DTM 120 
Lonicera sempervirens L.; DTM 402 
Sambucus nigra L.; DTM 1284 
Symphoriocarpus orbiculatus Moench; Raymond 865 


[riosteum angustifolium L.; DTM 2287 
Viburnum nudum L.; DTM 2286 
aay enPHE NCH jum; RDT 25303 
Raf.; MM 2823 


CRORE IEENGE RE 
naria benthamii Fenzl ex Torr.& A. Gray; RDT 65108 
LSU 

Arenaria lanuginosa (Michx.) oe DTM 1760 

*Arenaria sey llifolia L.; MM 261 

Cerastium brachypodum (Engelm.ex A. Gray) B.L. Rob- 
ins, DTM 2136 

*Cerastium fontanum Baumg.; Strout 122 LSUS 

*Cerastium salad Thuill.; DTM 937 

ackenzie sah 7284 

ee squarrosa suite Reid 4 

Minuartia drummondii aco McNeill; MM 7308 

Minuartia muscorum (Fassett) Rabeler; MM 7309 

Paronychia drummondii Tort. & A.Gray; DIM 2448 

Paronychia fastigiata (Raf.) Fern,; Correll & Correll 10096 
LSU 


Sagina decumbens (Ell.) Torr.& A. Gray; DTM 2723 


LAUUU PARISH, LOUISIANA 1209 


*Saponaria officinalis L, MM 237 
*Scleranthus annuus L.; a 82790 LSU 
Silene antirrhina L. ae 
*Silene gallica L; DTM 3 
*Silene Hociior Re a an 
Silene stellata (L.) Aiton f; DTM 2302 
Stellaria media (L.) Vill; Barbour 33 
*Vaccaria hispanica (P. Mill.) Rauschert; RDT 65131 LSU 


CELASTRACEAE 

vonymus americana L.; Hardy 8615 
CERATOPHYLLACEAE 
C lum d um L.; DTM 1554 
CHENOPODIACEAE 


iets ee (L.) AJ. Scott; RDT 65018 


Chenopodium berlandieri Moq.; RDT 66452 
*Che 


a 


Chenopodiu db.; RDT 65055 

Se um si sinnles (Torr.) Raf; DTM 935 

Chenopodium standleyanum Aellen; Overby 94 NLU 

Cycloloma atriplicifolium (Spreng.) Coult.,, Lewis 3663 
NLU 


Monolepis nuttalliana (JA. alana Greene; DTM 2196 
*Salsola tragus L RDT 65028 


CISTACEAE 

Helianthemum carolinianum (Walt.) Michx.; DTM 161 
Helianthemum georgianum Chapman; MM 2745 
Helianthemum rosmarinifolium Pursh; DTM 1744 
Lechea mucronata Raf.; MM 2715 

Lechea tenuifolia Michx.; DTM 1785 


CLUSIACEAE 


p reae (L.) Crantz; DTM 586 
Hypericum densiflorum Pursh; RDT 65084 LSU 
1ondii (Grev. & Hook.) Torr. & A. Gray; 


Hypericum drumn 
DIM 149 
1 frondosum Michx.; DTM 1257 
Byer icum pvaenuane ies: .) B.S.P; Allen 8539 LAF 
Hypericum gymnanthum Engelm. & A. Gray; RDT 
165816 


Hy peri 


um hypericoides (L.) Crantz; DTM 1 

Ay peictlen mutilum L.; ae 5317 

Hypericum nudiflorum Michx.ex Willd.; Raymond 1291 
LSUS 


a prolificum L. RDT 42445 LSU 

Hypericum pseudomaculatum Bush; DTM 453 
me cum mpunctotm Lam.; DTM 1066 

Triadenum walteri iJ. G.Gmel. ) Gleason; RDT 36265 


oct iadeladeiacaien 

*Col ulus arvensis L.; DTM 163 
Di cHonaia caroliniensis eee DTM 308 
Evolvulus sericeus Sw.; MM 3355 LSU 
!pomoea cordatotriloba Dennst.; DTM 696 
Ipomoea hederacea Jacq.; DTM 1920 


pomoea lacunosa L.; DTM 667 

pomoea panderata (L.) G.F.W.Mey.; DTM 1805 
pomoea purpurea (L.) Roth; MM 125 
pomees auamgel it L. RDT 42043 


*/pomoea wrightii Gray; DTM 2693 
Jacquemontia tamnifolia (L.) Griseb.,; DTM 640 
Stylisma humistrata (Walt.) Chapman; RDT 92409 
Stylisma pickeringii (Torr. ex M.A. Curtis) A. Gray; MM 
2677 


ee 
ornus drummondii C.A. Mey.; Hardy 8607 

zee florida L; Graham 571 LSUS 

Cornus foemina P. Mill. Le ee 

Nyssa aquatica L; RDT 3 

Nyssa eee ca Marsh.; ae 638 (includes WN. biflora T. 
Walte 


rigbenaminsane 

tica (L.) Schoenl.; RDT 34511 
Penthorurn sedoides _ ae 683 LSUS 
RDT 83128 NLU 


CUCURBITACEAE 

Cayaponia quinqueloba (Raf) Shinners;RDT 25353 LSU 

Melothria pendula L., DTM 233 

ee acene 

Cu Juss.ex Choisy; DTM 1934 

Cuscuta gronovi ii Willd. ex. J.A. pi MM 696 

Cuscuta indecora Choisy; MM 

Cuscuta pentagona Engelm.; ae 1788 

DROSERACEAE 

Drosera brevifolia Pursh; DTM 832 

EBENACEAE 

Diospyros virginiana ke DTM ] 176 

ELAEAGNACEAE 

*Elaeagnus pungens Thunb.; DTM 2054 (questionably 
naturalized 


aa 


ERICACEAE 

Lyonia mariana (L.) D.Don; RDT 33683 

Rhododendron canescens (Michx.) Sweet; RDT 33665 

Rhododendron oblongifolium (Small) Mallais; RDT 
88707 


d| osum (L.) Torr; DTM 2278 

Vaccinium ae Marsh.; DTM 1637 

Vaccinium corymbosum L.; Haynes 4628 (includes V. 
elliottii Chapman, V.fus n,and V.virgatum 
Aiton 

Vaccinium stamineum L.,; DTM 1155 


EUPHORBIACEAE 

fi at gracilens Gray; DTM 
calypha monococca jee aA Gray) L. Mill & Gan- 
dhi; ROT 

Acalypha es Riddell; DTM 1392 

Acalypha rhomboidea Raf.; DTM 704 

Acalypha setosa A. Rich.; DTM 1092 


atum Ait 


1210 


Acalypha virginica L., DTM 1371 
Chamaesyce cordifolia (Ell.) Small; DTM 1940 
Chamaesyce humistrata (Engelm.) Small; RDT 36205 
Chamaesyce maculata (L.) Small; DTM 1821 LSU 
Chamaesyce nutans (Lag.) Small; Haynes 4223 
Chamaesyce prostrata (Aiton) Small; DTM 1830 
Chamaesyce serpens (Kunth) Small; RDT 32893 
Cnidoscolus texanus (Muell.-Arg.) Small; Haynes 5414 
Croton argyranthemus Michx.; RDT 81395 LSU 
Croton capitatus Michx.; DTM 11 
Croton glandulosus L.; DTM 188 
Croton michauxii G.L.Webster; MM 2772 
Croton monanthogynus Michx,; DTM 170. 
Croton willdenowii G.L Webster; DTM 532 
aati corollata L.; DIM 71 

us dentata Michx.; MM 31 


72 
Euphorbia tetraspora Engelm.; Moore s.n. LSU 
A abt us carolini iensis Walt., MM 698 1 
aria L.; RDT 166975 
*Rick inus communis L.; Thieret 21037 LAF (questionably 
naturalized) 
sia lingia sylvatica Garden ex L.; DTM 1 
ragia eetonisioile Nutt; RDT 45419 
ichx hee 


TGUGLU INI 4" 


tears urticifol lia Michx.;D 45 
*Triadica sebiferum (L.) a a 1757 


FABACEAE 
Acacia angustissima (P. Mill.) Kuntze; ta 74 
*Albizia julibrissin Durazz., DTM 1 
pibllene fruticosa L.; MM 6793 

Amorpha paniculata Torr.& A. Gray; MM 255 
Amphicarpaea bracteata fe “ ae DTM 650 
Apios americana Medik. 
Astragalus canadensis ‘ ay an 
Astragalus distortus Torr.& A. Gray; DTM 260 


=> 


is soxmaniorum Lundell; DTM 1806 
Baptisia alba (L.) Vent.; DTM 867 
Baptisia bracteata Muhl. ex Ell; MM 176 
Baptisia nuttalliana Small; DTM ed 
Baptisia sphaerocarpa Nutt.; MM 1 
Centrosema virginianum (L.) pent a 169 
Cercis canadensis L.; Barbour 
silica fasciculata ae Greene; DTM 86 
maecrista nictitans (L.) on DTM 215 

arene mariana. DTM 1 
Crotalaria sagittalis L.; see 
a talaria fee ed Roth; ee z 
ex Willd; DTM 
ae phleoides a &A, ues eae MM 2657 
Dalea purpurea Vent; DTM 534 

Dalea villosa (Nutt.) Fie DIM 1797 


LAF 


AAD KAT 


BRIT.ORG/SIDA 22(2) 


Desmanthus illinoensis (Michx.) MacM.ex B.L. Robins. & 
Fern.; 1969 

(L.) DC; DTM 1973 
(Muhl. ex Willd.) DC.; DTM 211 
Desmodium glabellum (Michx.) DC.; RDT 37293 
pons ium hes old ee DC.; RDT 136869 

odium nudiflorum ( C.,;, Overby 292 NLU 
se Schub.; Raymond 2069 


[) di 


Desmoditim ni itt hij 


LSUS 


oditnm pan 


iculatum (L.) DC; Raymond 782 LSUS 
Desmodium sessloliam (Torr) Torr.&A. se DIM 1084 
Desmodium viridiflorum (L.) DC.; RDT 3 

Dioclea multiflora (Torr. & A. Gray) C. a 3729] 
Sna herbacea L., DTM 439 


Galactia volubilis (L.) Britt; DTM 591 

Gleditsia ia. aquati ica Marsh,; Reid 4636 NLU 
leditsia triacanthos L.; Hardy 8612 

Glottidium vesicarium Jacq.) Harper; Haynes 5328 
eu striata (Thunb.) Schindl.; DTM 1505 


L.;DITM 37 
bans pusillus Ell; DTM 795 
Lathyrus venosus Muhl. ex Willd.; DTM 2199 
*Lespedeza cuneata (Dum.-Cours.) G. Don; RDT 36216 
Lespedeza hirta (L.) Hornem.; DTM 678 
Lespedeza procumbens Michx.; DTM 733 
Lespedeza repens (L.) W. Bart.; re 
Lespedeza stuevei Nutt.; MM 2 
eae a violacea (L.) Pers.; aa i 


le 


inica (L.) Britt., DTM 
ae uni oliolatis (Hook.) Benth.; ae 66462 LSU 
Be Inus texensis ae Newsom s.n. LSU 
(L.) Huds.; ae 362 
ona) cago ating L; DTM 1 
(L.) L. RDT oe 
“Media polymorpha L.; oo 530 
Medicago sativa L.; DTM 1 
*Melilotus albus Medik.; ane 
*Melilotus indicus (L.) All, DTM “ 
*Melilotus officinalis (L.) Lam. DTM 410 


Mimosa nuttallii (DC.) B.L. Turner: “it 164 


ee 


Mimosa Strigillosa Torr.& A. Gray; DIM 19 
Neptunia lutea ese atek ie ;DTM 50 
Neptunia pubescen h.; Lasseigne 1002 LAF 


Orbexilum pene (P. Mill.) Rydb,; DTM 385 

Orbexilum simplex (Nutt. ex Torr.& A. Gray) Rydb,; DTM 
1119 

Parkinsonia aculeata L.; DTM 756 

Pediomelum digitatum (Nutt. ex Torr. & A. Gray) Isely; 
MM 235 


Pediomelum hypogaeum (Nutt.ex Torr.& A.Gray) Rydb,; 
M 


M 2549 
Prosopis glandulosa Torr; Cocks s.n. NO 
*Pueraria montana (Lour.) Merr.; Hardy 8896 
Rhynchosia latifolia Nutt. ex Torr.& A. Gray; DTM 520 
Rhynchosia minima (L.) DC; MM 6991 
Rhynchosia reniformis DC.; DTM 129 


LAUUY PARISH, LOUISIANA 1211 


Rhynchosia tomen t (L.) Hook. & Arn.; RDT 36931 
ene isola i MM ia 
Robinia pseudoacacia L.; Haynes 5699 


Senna iene (L.) Link; DTM 196 

Senna obtusifolia (L.) Irwin & Barneby; DTM 1575 
Sesbania drummondii (Rydb.) Cory; DTM 202 
Sesbania herbacea (P. Mill.) McVaugh; DTM 1476 
*Sesbania punicea (Cav.) Benth.; DTM 1215 

Sophora affinis Torr.& A. Gray; Cocks s.n. NO 
Strophostyles helvula (L.) Ell; DIM 564 

Strophostyles leiosperma (Torr.& A. Gray) Piper; DTM 544 
Strophostyles umbellata (Muhl.ex Willd.) Britt; Raymond 


SUS 
Stylosanthes biflora (L.) B.S.P; MM 2713 


ee brychoides Nutt.; DIM 1904 
Te phrosia Migtolaaa (lis) Pers.; ‘DTM 46 
rifolium a Seek 80865 
*Trifolium campestre Schreb.; DTM 151 
Trifolium carolinianum Michx.; DTM 2639 


*Trifolium dubium eee a 289 
¥Trifolitsm Incarnatiim L.;: DIM 


Trifolium reflexum L.; in 2638 
*Trifolium repens L.; DTM 136 
*Trifoliun upinatum L., DTM 313 
* Trifolium ee Savi; au 125 
Vicia caroliniana Walt.; DTM 2137 
Vicia ludoviciana bees a 1160 
*Vicia lutea L.; RDT 1 
Vicia anne FG. ies MM 152 
*Vicia sativa L.; 5 
*Vicia ee (li ) Schreb.; RDT 88360 LSU 
*VICIA villosa Roth; 394 
Wisteria frutescens (L.) Poir; Haynes 5356 
*Wisteria sinensis (Sims) DC.; DTM 1131 
Zornia bracteata J.F.Gmel.; MM 2844 


— 


FAGACEAE 
Castanea pumila (L.) P. Mill; DTM 1202 
Fagus grandifolia Ehrh.; ne 208 LTU 
Quercus alba L.; Barbour 56 
Sie qusagsaie ae MM 2872 
Michx.; Haynes 4094 
eee hemispherica Bart. ex Willd.; RDT 32931 
aU incana Bat ne: MM 581 
ee laurifolia Michx.; RDT 37236 
Quercus lyrata Walt; RDT 42015 
uercus i iaiaicelle ices: se nae LAF 
Qu ex Small; RDT 167037 
Quercus manleaaee Mu enh. Foymone 1321 LSUS 
Quercus michauxii Nutt.; DTM 
Quercus Se eae ot 36212 
Quercus nigra L.;M Tie 
Quercus phellos L.; eee 1292 LSUS 
Quercus rubra L.; Middlebrooks 206 NLU 
Quercus shumardii Buckl.; Hardy 8609 


Quercus stellata Wangenh.; MM 5016 (includes Quercus 
imilis Ashe) 


Quercus velutina Lam. Raymond 145 NLU 

Quercus virginiana P.Mill.; Rowe 105 LSUS 
FUMARIACEAE 

Corydalis flavula (Raf.) DC.; RDT 88228 

Corydalis micrantha (Engelm. ex A. Gray) A. Gray; MM 


L 00 ‘ | \A/\AS 


GENTIANACEAE 
Centauri a W.Wright ex Piper; 


MM 6435. 
Sabatia angularis (L.) Pursh; DTM 145 
Sabati pestris Nutt; DTM 72 
GERANIACEAE 
anium carolinianum L.; DTM 301 
mL;DTM 990 


GROSSULARIACEAE 
tea virginica L.; DTM 1196 
Ribes curvatum Small; DTM 2129 


HALORAGACEAE 

*Myriophyllum aquaticum (Vell.) Verdc.; Barbour 454 
Myriophyllum heterophyllum Michx.; a 908 
*Myriophyllum spicatum L.; Barbou 

Proserpinaca palustris L.; MM 205 


HAMAMELIDACEAE 
Hamamelis virginiana Ly ; Manning 1 176 LTU 
Liquidambar he L;DTM 1071 


HIPPOCASTANACEA 
Aesculus pavia L.; Baas 446 LSUS 


HYDROPHYLLACEAE 
Hydrolea ovata Nutt. ex ae DIM 12 
Hydrolea uniflora Raf.; D 
Phacelia glabra Nutt.; a - 
Phacelia strictiflora (Engelm.& A.Gray) A.Gray; MM 2544 


JUGLANDACEAE 

ee lba(L -) Nutt. ex Ell; Raymond ee mi 
quatica (Michx. f) Nutt; Barbour 

cane cor formis (Wangenh.) K. Koch; ee 1302 


* raninnm diccect 


a. ia (P. Mill.) Sweet; RDT 170092 

Carya illinoiensis (Wangenh,) K. Koch; MM 6470 
ee f) ail 503 LSUS 
ane ovata (P. Mill Koch; MM 

Carya texana Biel: “ rdy 8611 

Juglans nigra L.; Raymond 1679 NLU 


LAMIACEAE 
*Ajuga reptans L.; MM 202 

*Glechoma hederacea L; RDT 43545 
one Al eaee cain DIM 873 
L;DIM 53 
*Lamium purpureurn Ls DIM 2095 
Muhl. ex W. Bart; DTM 1 


7 


1212 


h; DTM 682 

Lyc Opus ViIrginicus Lee -RDT soi 
*Mentha spicata L.; DTM 1 
Monarda citriodora Cerv. ex e RDT 45372 
Monarda clinopodioides Gray; DTM 1753 
Monarda fistulosa L.; DTM 93 
Monarda punctataL.; DTM 15 
*Perilla frutescens (L.) Britt.;, MM 270 
Physostegia angustifolia Fern.; 2 - 

Physostegia digitalis Small; DTM 


Ph ial intermedia (Nutt.) in & A.Gray; DTM 


a is vulgaris L.; DTM 3 

Pycnanthemum albescens aa & A. Gray; DTM 194 

Enea RIC n (Michx.) Pers.; iat 6/061 
h M5 


Pycnanthemur PLCRIUTIUilUreil 
Salvia azurea Which ex Lam.; DTM 214 
Salvia lyrata L.; DTM 35 
Seutellaria cardiophylla ceae : A.Gray; MM 2723 
mained rea tiel ial; DIM4 

Hill; DTM . 
san laria ie es ar a 323 
Bet floridana se ie ex «Benth MM 214 
sae sila we DTM 
L.;DTM i 
L.; DTM 653 


Trick t Ait 


LAURACEAE 
Lindera kl eee Hardy 8617 
Sassafras albidum (Nutt.) Nees; DTM 1068 


LENTIBULARIACEAE 

Uticularia gibba .; Barbo eae 

Uticularia inflata Walt; DTM 2 

Uticularia macrorhiza sees len 464 
LINACEAE 

Linum medium (Planch.) Britt.; DTM 522 
‘inum striatum Walt.; RDT 170116 
LOGANIACEAE 

Gelsernium sempervirens (L.) Aiton f; DTM 270 


j—~< 


Mitreola petiolata (J.F. mel.) Torr. & A. Gray; DTM 201 


Spigelia marilandica (L.) L.;MM 210 


LY THRACEAE 

mmannia auriculata Willd.; ra . 7 
Ammannia coccinea Rottb.; M 
orca te us (L.) Ell; sis on LSUS 
Lythrum alatum Pursh ; MM 6783 


Rotalo ra 


ior (L.) | Koeht 1e; f laynes 5336 


MAGNOLIACEAE 
Magnolia grandiflora L.; Burns 238 LSUS 
Magnolia virginiana L.; Barbour 1191 


MALVACEAE 
*Abutilon theophrasti Medik.; RDT 119031 


BRIT.ORG/SIDA 22(2) 


Callirhoe alcaeaides (Michx.) A. ay DTM 1049 
a rhoe digitata Nutt; RDT 8837 
allirhoe involucrata (Torr. & an A. Gray; Newsom 


Callirhoe papaver (Cav.) A. a MM 257 
;DIM9 


Hibiscus laevis Al 


Hibiscus moscheutos L.; oe 168 

*Hibiscus syriacus L.; Gamblin s.n. LTU (localized and 
ersisting) 

Malvaviscus arboreus Dill.ex Cav.; MM 123 

Modiola caroliniana (L.) G. Don; DTM 379 

Sida rhombifolia L.; DTM 186 

Sida spinosa L.; DTM 521 


MELASTOMATACEAE 

Rhexia mariana L.; DTM 243 
MELIACEAE 

*Melia azedarach L.; MM 6183 

sates a balsas 

(L.) DC; Barbour 473 


MOT ene ene 

Mollugo verticillata L.; DTM 1373 
MONOTROPACEA 

Monotropa ica L.; Thieret 24524 LAF 

Monotropa uniflora L.; MM 281 


MORACEAE 

*Broussonetia papyrifera (L.) UHer.ex Vent. DTM 1145 
Maclura ae (Raf.) aoe Barbour 570 
*Morus 

Morus a L.; Haynes iD 


MYRICACEAE 
Morella caroliniensis (P. Mill) Small; RDT 33652 
Morella cerifera (L.) Small: Haynes 4507 


NELUMBON 
Nelumbo lutea ie Barbour 467 


NYCTAGINACEAE 
Boerhaavia rite L. DTM 2573 
ete ia abe L.; RDT 65066 LSU 
da (Walt.) Heimerl; MM 2610 


NYMPHAEACEAE 
Nuphar advena (Aiton) W.T. Aiton; MM 208 
Nymphaea odorata Aiton; DTM 9 


OLEACEAE 

Chionanthus virginicus L.; Raymond 1307 LSUS 

Forestiera acuminata (Michx.) Poir; Barbour 1004 

Forestiera clan ee Poir.; Raymond s.n.LSUS 
axinus analjH 613 

Fraxinus a P. vey M252 

Fraxinus pennsylvanica Marsh.; MM 6682 

*Ligustrum japonicum Thunb.; DTM 1173 

*Ligustrum lucidum Aiton f, RDT 32907 

*Ligustrum sinense Lour; DIM 1171 


a 


CADDO PARISH, LOUISIANA 1213 


ONAGRACEAE 

Gaura longiflora Spach; DTM 221 

Gaura mollis James; DTM 104 

Gaura sinuata Nutt. ex Ser.; DTM 1790 

Ludwigia alternifolia L, DTM 115 

Ludwigia decurrens Walt., DTM 617 

non a aa Walt.; DTM 1400 

Ludwigia hi .; DTM 1301 

Ludwigia leptocarpa ses DTM 226 
Ludwigia linearis Walt; 

Ludwigia palustris (L.) i oe 4090 

Ludwigia peploides (Ku oy hee Bes 4087 
Ludwigia repens J.R. For 

Oenothera biennis L.; ie 

Oenothera heterophylla een aia 31220 LAF 
Oenothera laciniata Hill; DTM 7 

Oenothera linifolia Nutt, DTM : ] 

Oenothera spachiana Torr. & A.Gray; DIM 989 
Oenothera speciosa Nutt; DTM 122 


OXALIDACEAE 
Oxalis corniculata L., MM 5947 


*Oxalis rubra St.-Hil. DTM 492 
Oxalis violacea L., DIM 417 


PAPAVERACEAE 


PASSIFLORACEAE 
Passiflora incarnata L; DTM 133 
Passiflora lutea L.; DTM 96 


PEDALIACEAE 

Proboscidea louisianica (P. Mill.) Thellung; Hall s.n.LSUS 
PHRYMACEAE 

Phryma oa DTM 1290 

PHYTOLACCA 


Phytolacca ie L.; DTM 61 
Rivinia humilis L.; DTM 1490 


PLANTAGINACEA 
ey DTM 1025 


Plantago hookeriana Fisch. & C. = Mey.; MM 2580 
*Plantago ignecolaea DTM 8 
Plantago | fT see 
Plantago pusilla Nutt; 7 730 

Plantago anes oe Dene; a 65015 
ica L.; DTM 1026 

na ADeneE DTM 913 


i] - | 


D 
FICOge virgin 


re a MUGY VE ightia 
PLATANACEAE 
Platanus occidentalis L.; Haynes 4634 
POLEMONIACEAE 

Phlox divaricata L.; Lawrence 17 LTU 
Phlox drummondii Hook.; MM 197 


Phlox pilosa L.; Raymond 1200 LSUS 


POLYGALACEAE 

Palygala mariana P. Mill; RDT 170119 

ke a polygama Walt; MM 2629 
olygala sanguinea L., DTM 461 

Polygala verticillata L., RDT 36193 


POLYGONACEAE 

Brunnichia ovata (Walt.) Shinners; DTM 171 
Eriogonum longifolium Nutt; MM 2323 

Eriogonum multiflorum Benth.; MM 2870 

Polygonella americana (Fisch. & C.A. Mey.) Small; MM 


aly 


lygonum avi DTM 1336 
*Polgorum ey Blume; RDT 170115 
, RDT 66461 LSU 
Polygonum densiflorum Meisn.; DTM 608 


L; RDT 32876 


ee hydropiperoides Michx.; DTM 113 
*Polygonum lapathifolium L., DTM 601 
Polygonum pensylvanicum L., DTM 213 
*Polygonum persicaria L.; Haynes 4222 
Polygonum punctatum Ell; Haynes 4224 
Polygonum ramosissimum Michx.; DTM 643 
Polygonum scandens L.; MM 292 

ee setaceum Baldw.; DTM 239 

nN virginianum fs a 775 SUS 
pare altss imus Wood; 

*Rumex crispus L.; DTM 81 - 

Rumex hastatulus Baldw.; DTM 798 

*Rumex pulcher L.; DTM 865 

Rumex verticillatus L.; DTM 907 


PORTULACACEAE 

| Uyltr nia virginica L.; ‘DIM 11 
Portulaca oleracea L.; DTM a 
Talinum parviflorum Nutt, MM 3354 
Talinum rugospermum Holz.; MM 2757 
PRIMULACEAE 
*Anagallis arvensis L.; Lynch 3861 LSUS 
ana minima (L.) Krause; a ies 


dec-atheon meadia | 


Lysimachia lanceolata Walt.; ee 565 LSUS 
Samolus valerandi L., DTM 374 


RANUNCU 

Anemone reste Pritz., DIM 2590 

Anemone caroliniana ae DTM 264 

Clematis crispa L.; MM 2 

Clematis reticulata Walt.; — 2680 

*Clematis terniflora DC,; ey - 

Clematis virginia Chih lee MM 2 

Delphinium carolinian um ae MM 2774 
Myosurus minimus L.; Barbour 556 

Ranunculus re L. DTM 953 

Ranunculus fascicularis Muhl.ex Bigelow; DTM 773 
Ranunculus laxicaulis (Torr.& A. Gray) Darby; DTM 292 


1214 


*Ranunculus marginatus d'Urv,; DTM 980 
*Ranunculus muricatus L.; DTM 409 
*Ranunculus parviflorus L.; DIM 878 
Ranunculus pusillus Poir; OTM 775 
urvatus Poir; DTM 1047 
*Ranunculus sardous Crantz; DTM 1048 
Ranunculus sceleratus L.; DTM 2019 

Ee ce trilobus Desf; DTM 902 

da m Fisch. & Ave-Lall.; DIM 901 


RHAMNACEAE 

Berchemia scandens (Hill) K. Koch; DTM 1781 
Ceanothus americanus L.; MM 2370 

Ceanothus herbaceus Raf; RDT 93612 

Frangula caroliniana (Walt.) A.Gray; Hardy 8646 

*Ziziphus zizyphus (L.) Karst. RDT 33078 

ROSACEAE 

Agrimonia microcarpa Wallr.; Haynes 5337 

Agrimonia rostellata Wallr.; Haynes 5315 

Amelanchier arborea ( Michx.f.) Fernald; Manning 485 


Ranunculus re 


LTU 
*Aphanes microcarpa (Boiss. & Reut.) Rothm.; RDT 


82 
Aronia arbutifolia (L.) Ell; RDT 33632 
Crataegus berberifolia Torr.& A.Gray; Graham 460 LSUS 
Crataegus brachyacantha Sarg. & Engelm.; Rowe 85 
LSUS 


Crataegus crus-galli L.; RDT 165825 

Crataegus engelmannii Sarg.; DIM 1966 LSU 
ig ee marshallii Egglest.; Haynes 4623 
Crataegus opaca Hook. & Arn.; Haynes 4627 
Caton apatiiiata Michx.; Haynes 4095 


hh. MM 2578 


sai viridis L.; Barbour 1164 
*Duchesnea indica uae: foe DIM4 
in Ansty, ~ 


Malus canes (Aiton) vee 88278 
Porteranthus stipulatus (Muhl.ex le Britt.; DTM 1030 
*Potentilla recta L.; MM 2620 
Potentiilla simplex Michx., DTM 48 
Prunus americana Marsh.; RDT 96045 
Prunus angustifolia Marsh.; MM 2538 
Prunus caroliniana (P. Mill.) Aiton; DTM 1121 
Prunus gracilis Engelm.& A. Gray; MM 2539 
Prunus mexicana S.Wats.; Raymond 69 LSUS 
Prunus serotina Ehrh.; Barbour 47 
Prunus umbellata Ell; DTM 1603 
M is 


1.p A rd . | 
' 


*Rosa bracteata J.C.W 
naturalized) 

Rosa carolina L.; MM 201 

*Rosa laevigata Michx.; Barkley 103 LSU (questionably 
naturalize 

*Rosa multiflora Thunb. ex Murr; Lynch 2273 LSUS 
(questionably naturalized) 


BRIT.ORG/SIDA 22(2) 


Rosa setigera Michx.; MM s.n. 
Rubus argutus Link.; Raymond 2006 LSUS 
Rubus trivialis Michx.; Graham 444 LSUS 


RUBIACEAE 
Cephalanthus Heater is DTM 1296 
Diodia teres Walt. 
di Ua virginiana las one _ 
Gah ium pee Le OT) 288 
ful ans Michx.; DTM 922 
Galiu ium abuse Bigelow; fa 994 
*Galium parisiense L.; 323 
Galium pilosum Aiton; DTM 915 


Galium triflorum Michx.; RDT geen 
Galium uniflorum Michx.; RDT 
Hedyotis nigricans (Lam.) ase ae 227 
oustonia micrantha (Skinner) Terrell; DTM 1979 
Houstonia pusilla Schoepf; Raymond 741 LSUS 
Houstonia rosea (Raf.) Terrell; DTM 255 
Mitchella repens L.. DTM 378 
Oldenlandia boscii (DC.) Chapman; DTM 2996 
Oldenlandia uniflora L.; ie ae 
nna scabra L T 


ans nee 9 
Spermacoce glabia Michie! DIM 1 


RUTACEAE 
*Poncirus trifoliata (L.) R 
a oie ata L.; RD 


ava ae if Raymond 13 


a _ 


19LSUS 
SALICACEA 

*Populus Ae L.; DIM 2230 

Populus deltoides Bartr.ex Marsh.; Haynes 4633 
Salix exigua Nutt; Haynes 4157 

Salix humilis Marsh. RDT 88359 LSU 

Salix nigra Marsh.; Haynes 5700 

SAPINDACEAE 

Cardiospermum halicacabum L.; Haynes 5292 
Sapindus saponaria L.; RDT 88240 

riniaanates 


1Michx.; MM 2828 
. RDT 33011 


Sideroxvlon Ivcioides L 


SAURURACEAE 
Saururus cernuus L.,; DTM 30 


SAXIFRAGACEAE 
Heuchera americana L.; Hay 
ea eke ies ai . 34504 


SCROPHULARIACEAE 

Agalinis ae Pennell; Pennell 5658 PH 
Agalinis fasciculata (Ell.) Raf; DIM 755 

Agalinis gattingeri (Small) Small; DTM 541 

Agalini is heterophylla (Nutt.) ex Britt; DIM 736 
Agalinis homalantha Pennell; DTM 748 

Agalinis purpurea (L.) Pennell; a 2570 


BERT. AAVUY TART, LOUISIANA 1215 


Agalinis tenuifolia (Vahl) Raf; ate a 

Ag alinis Vil idis (Small) Penn ell: D 

Aureolaria grandiflora (Benth.) et DTM 605 
Aureolaria pectinata (Nutt.) Pennell; Cocks s.n.NO 
Bacon g Kk poungiols en) Wettst., DTM 1592 


aie indivisa ee Barbour 
*Chaenorhinum minus (L.) Lange; ae 65115 LSU 
Collinsia violacea Nutt.; Maynard 151 LTU 
Dasistoma macrophylla (Nutt.) Raf; Palmer 10611 MO 
Gratiola flava Leavenworth; MM 7285 
Gratiola neglecta Torr; DTM 398 
Gratiola pil osd Michx.; RDT 33636 
Gratiola virginiana L.; DTM 1127 
ee multifida (Michx.) Nutt.; DTM 1103 
Lindernia crustacea (L.) F. Muell.; RDT 41803 
Lindernia dubia (L.) Pennell; DTM 165 
*Mazus pumilus (Burm.f) Steenis; DTM 964 
Mecardonia acuminata (Walt.) Small; DTM 216 
Mimulus alatus Aiton; DTM 603 
Nuttallanthus canadensis (L.) D.A. Sutton; DTM 263 
Nuttallanthus texanus (Scheele) D.A. Sutton; DTM 381 
*Pqrentucellia viscosa (L) Caruel; MM 2627 
Beale ial is ae s Ly DTM 37 
is Nutt. ex Sims; ao 554 LSUS 
Penstemon iain Pennell; DTM 
Penstemon murrayanus ee MM o 7 
Penstemon tenuis Small M557 
Penstemon tubaeflorus ce DTM 81 
Scrophularia marilandica L; RDT 88232 
*Verbascum blatteria L. Hardy 6 
*Verbascum thapsus L.; DTM 200 
*Veronica arvensis L.. DTM 262 
*Veronica hederifolia L; Burns 13 LSUS 
Veronica peregrina L.; DTM 303 
*Veronica persica Poir.; MM 5966 
*Ver UT Ca polita FI ies; DIM 273 


SIMAROUBACEAE 
*Ajlanthus altissima (P. Mill.) Swingle; Lewis 2185 NLU 


SOLANACE 
fee stramonium L pie 930 
Watd |: 

Physalis cineraesens (Dunal) ; S.Hitche,; RDT 32855 

Physalis cordata P. Mill; RDT 32854 

ede hederi i ia Gray; RDT 89017 
salis f [| Nees; MM 2619 

Physalis longifol ia Nutt. MM 6667 

Physalis mollis Nutt.; MM 2608 

Physalis pubescens L; DIM 1779 

oe pu) Nutt; RDT 140326 


eL.; DIM 99 


Gc lanum i fj 


R af.; Petersen s.n.LSU 
Solanum elaeagnifolium Cav.; DTM 108 
*Solanum pseudocapsicum L.; Raymond 2149 LSUS 
hanthum Dunal; DTM 118 


Solanum nf\ 
| ia 


Salanum rostratum Dunal: RDT 81358 
SPHENOCLEACE 

*Sphenoclea aoa ee Gaertn.; RDT 36943 
STYRACACEAE 

Styrax americanus Lam.; MM 189 

Styrax grandifolius Aiton; RDT 37239 
SYMPLOCACEAE 

Symplocos tinctoria (L.) Her; Barbour 1166 
TAMARICACEAE 

*Tamarix gallica L, DTM 1318 

TILIACEAE 

Tilia americana L; DTM 1144 


— 


= 


ULMACEAE 

Celtis laevigata Willd.; Haynes 4092 
Celtis tenuifolia Nutt.; RDT 37283 
Planera aquatica J.F. Gmel.; Haynes 8648 
ie alata Michx.; Haynes 4508 

us americana L. Haynes 4574 
ae crassifolia Nutt.; Hardy 8610 

Ulmus rubra Muhl.; Raymond 308 LSUS 


URTICACEAE 

Boehmeria cylindrica (L.) Sw, DTM 224 

Parietaria pensylvanica Ae ex a DIM 1796 
Pilea pumila (L.) A. Gray; M 

Urtica chamaedryoides ee oa 3 


VALERIANACEAE 
Valerianella radiata (L.) Dufr; DTM 1028 


VERBENACEAE 

Callicarpa americana L; DTM 1 

Glandularia bipinnatifida (Nutt. ; ie RDT 99514 
Glandularia canadensis (L.) Nutt; DTM 9 

*Glandularia pulchella (Sweet) Troncoso; DTM 1355 

*Lantana camara L; RDT 36934 

Lantana urticoides Hayek; DTM 1570 

Phyla lanceolata (Michx.) Greene; DTM 22 

Phyla nodiflora (L .) Greene; DTM 9 

*Verb 


5 


Verbena nate I; D 

I a 893 

Verbena na [ss DTM 
440 


Verbena xutha Leh RRs 


*Verbel 1A 


*Vitex agnus-castus . MM 226 


VIOLACEAE 

Viola bicolor Pursh; DTM 257 

Viola lanceolata L., DTM 2106 

Viola palmata L; DTM 772 

Viola pedata L., DTM 43 

Viola primulifolia L.; DTM 21 a 

He gee pus RDT 1 
iola DTM aoe 


la sagittata Aito 


1216 BRIT.ORG/SIDA 22(2) 


Viola sororia Willd.; DTM 788 Parthe issus quinquefolia (L.) Planch.; MM 6208 
Viola villosa Walt. MM 141 Vitis aestivalis Michx., DTM 912 
Viola walteri House; DTM 816 Vitis cinerea (Engelm.) Millard; Correll & Correll 10091 
VISCACEAE Prati ane 
Phoradendron tomentosum (DC) Engelm. ex A. Gray; vitis palmata es RDT 25315 

DTM 2102 Vitis riparia Michx., DTM 1159 


Vitis rotundifolia Michx.; MM 6201 
VITACEAE Vitis vulpina L.; RDT 42042 
Ampelopsis arborea (L.) Koehne; DTM 152 
Ampelopsis cordata Michx., DTM 905 ZYGOPHYLLACEAE 
Cissus incisa Des Moul.; DTM 1784 *Tribulus terrestris L., DTM 600 


ACKNOWLEDGMENTS 


Stephanie Aamodt, former Chairman of the Biology Department, Louisiana State Uni- 
versity in Shreveport, and Al Vekovius, former Dean of Science, Louisiana State Univer- 
sity in Shreveport, provided the space for an herbarium where the work could be done. 
Guy Nesom, BRIT; Larry Brown, SBSC; Billie Turner, TEX, and Diane Ferguson, LSU, helped 
in determining several problematic specimens. Diane Ferguson, LSU; Kerry Heafner, NLU; 
Garrie Landry, LAF; Amanda Neill, BRIT; Eric Sundell, UAM; Milan Vavrick, LTU: Anne 
Bradburn, NO; Sherri Herndon, DUKE: George Russell, US,and James Solomon, MO, loaned 
specimens. Thanks also to Chris Reid, Louisiana Natural Heritage Program; Charles Allen, 
Botanist, Fort Polk; the late John Thieret, KNK; Larry Raymond, Caddo Parish Parks De- 
partment, and Gary Hanson, Patricia Peyton, and Scott Foord of the Red River Watershed 
Management Institute, Louisiana State University-Shreveport. Paul Sisson, Dean of Sci- 
ence, Louisiana State University in Shreveport, obtained funds for publication costs. 
Charles Allen, R.D.Thomas, and an anonymous reviewer provided helpful comments on 
the manuscript. 


REFERENCES 

Aiten,C.M.,D.A. Newman, and H. Winters. 2004. Grasses of Louisiana. 3rd Edition. Allen’s Native Ventures, 
Pitkin, Louisiana. 

Braun, E.L. 1950. Deciduous forests of eastern North America. Hafner Press, New York. 

Canne-Hivuker, JM. and M. Dusrute. 1993. A new species of Agalinis (Scrophulariaceae) from Grimes 
County, Texas. Sida 15:425-440, 

Corrett, D.S. and H.B. Corrtit. 1941. A collection of plants from Louisiana. Amer. MidI. Naturalist 26: 
30-64. 

Diccs, G.M., B.L. Lipscome, and R.J.O’Kennon, 1999. Illustrated flora of north central Texas. Sida Bot. Misc. 
16:1-1626, 

Diccs, G.M., B.L. Lirscome, M.D. Reep, and RJ. O’Kennon. 2006. Illustrated flora of east Texas. Sida Bot Misc. 
26:1-1594., 

Epwarbs, J.P, P.G. Martin, J.W. Macoun, W.W. Kicpatrick, and C. Henry. 1980. Soil survey of Caddo Parish, 
Louisiana. U.S.D.A. Soil Conservation Service, Washington, D.C. 

Flora or Nort America Eptroriat Comirtee (eds.). 1993-2006. Flora of North America North of Mexico. 
Vols. 1, 2, 3,4, 22, 23, 25, 26. Oxford Univ. Press, New York and Oxford. 

Fiores, D.L. 1984. Jefferson and southwestern exploration: The Freeman and Custis accounts of the 
Red River expedition of 1806. Univ. Oklahoma Press, Norman. 

GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida Bot. Misc. 4:1—202. 


Be Ah ace RISH, LOUISIANA 1217 


Karresz, J.A. and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina 
Botanical Garden. Chapel Hill. 

Ker, E.L,, JR. SincHurst, and S. Cook. 2004, Geocarpon minimum (Caryopyllaceae), new to Texas. Sida 
21:1165-1169. 

Larke, J.O.and L.M. Switn. 1994. Rare plants of pine-hardwood forests in Louisiana. Louisiana Depart- 
ment of Wildlife and Fisheries, Baton Rouge. 

Leipotr, A., S.McDaniet, and T. Nuttie. 2002. The flora of Oktibbeha County, Mississippi. Sida 20:691—765. 

Lester, G.D,, S.G. Sorensen, PL. Fauckner, C.S. Reio, and .E. Maxi. 2005. Louisiana Comprehensive Wildlife 
Conservation Survey. Unpublished report. Louisiana Natural Heritage Program, Louisiana Depart- 
ment of Wildlife and Fisheries, Baton Rouge. 

Louisiana Naturat Heritace Procram. 2004. Rare plant species of Louisiana. Unpublished report. Louisi- 
ana Natural Heritage Program, Louisiana Department of Wildlife and Fisheries, Baton Rouge. 
Louisiana Naturat Heritace Procram. 2006. Rare plant species of Louisiana. Unpublished report. Louisi- 
ana Natural Heritage Program, Louisiana Department of Wildlife and Fisheries, Baton Rouge. 
MacRoseats, B.R. and M.H. MacRoserts. 1994. Floristics of a xeric sandyland in western Louisiana. 

Phytologia 77:414-424,. 

MacRoseats, B.R.and M.H.MacRoserts. 1995. Vascular flora of xeric sandhills in northwestern Louisiana. 
Phytologia 79:123-131. 

MacRoserts, B.R. and M.H. MacRoserts. 1996. Floristics of xeric sandhills in east Texas. Phytologia 80: 
desk. 

MacRoserts, B.R. and M.H. MacRoserts. 1997. Talinum rugospermum Holz. new to Louisiana with notes 
on terete-leaved Talinum in Louisiana. Phytologia 82:86-93. 

MacRoserts, B.R., MH. MacRoserts, and J.C. CatHey. 2002. Floristics of xeric sandylands in the post oak 
savanna region of east Texas. Sida 20:373-386. 

MacRoeerts, D.T. 1977a. Additions to the Louisiana flora. Sida 7:220-222. 

MacRoseats, D.T. 1977b. New combinations in Tradescantia. Phytologia 37:451-452. 

MacRoserts, D.T. 1978a. Notes on Tradescantia: T. diffusa Bush and T. pedicillata Celarier. Phytologia 
38:227-228. 

MacRoserts, D.T. 1978b.The status of Agalinis caddoensis Pennell. Phytologia 40:1-6. 

MacRoserts, D.T. 1979a. Checklist ifthe plants of Caddo Parish, Louisiana. Bull. Mus. Life Sci,, Louisiana 
State University-Shreveport 1:1-54. 

MacRoserts, D.T. 1979b. Notes on Tradescantia Ill. Tradescantia ohiensis Raf. var. paludosa (Anderson 
and Woodson) MacRoberts, comb. nov. Phytologia 43:380-382 

MacRoserts, D.T. 1980a. Notes on Tradescantia IV: The distinction between Tradescantia virginiana 
(Commelinaceae) and Tradescantia hirsutiflora. Phytologia 46:409-416. 

MacRoserts, D.T.1980b.Notes on Tradescantia (Commelinaceae) V. Tradescantia of Louisiana. Bull. Mus 
Life Sci., Louisiana State University-Shreveport 4:1—-15. 

MacRoserts, D.T. 1980c. Notes on Tradescantia VI. Observations on the mechanism of pigment depo- 
sition. Phytologia 46:41 7-418. 

MacRoserts, D.T. 1984a.The vascular plants of Louisiana. An annotated checklist and bibliography of 
the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci., Louisiana 
State University-Shreveport 6:1-165. 

MacRoserts, 0.1.1984b.Helianthemum rosmarinifolium and Oenothera triloba in Louisiana. Phytologia 
55:343. 

MacRoeerrs, D.T. 1987.Reported Texas endemics. Castanea 52:72. 


1218 BRIT.ORG/SIDA 22(2) 


MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. 
Life Sci., Louisiana State University-Shreveport 7—9:1-756. 

MacRoserts, D.T.,B.R.MacRoserts, and M.H.MacRostrts. 1997.A floristic and ecological interpretation of 
the Freeman and Custis Red River Expedition of 1806. Bull. Mus. Life Sci., Louisiana State Univer- 
sity-Shreveport 12:1-26. 

MacRoserts, M.H. and B.R. MacRoseats. 1998. Quality change of habitat in northwestern Louisiana. 
Phytologia 84:297-303. 

MacRoseets, M.H., B.R. MacRoserts, B. Sorrie, and R. Evans. 2002. Endemism in the West Gulf Coastal 
Plain: importance of xeric habitats. Sida 20:767-779. 

MacRoberts, M.H., B. R. MacRoserts, and L.S. Jackson. 2003. Louisiana prairies. In: E. Peacock and T. 
Schauwecker, eds. Blackland Prairies of the Gulf Coastal Plain: Nature, Culture and Sustainability. 
Univ. Alabama Press, Tuscaloosa. Pp. 80-93. 

MacRoserts, M.H. and B.R. MacRoserts. 2003a.West Gulf Coastal Plain ecoregions. Sida 20:1245-1274. 

MacRoserts, M.H.and B.R.MacRoserts.2003b.The east-west transition of flora in Texas:a biogeographical 
analysis. Sida 20:1693-1700. 

MacRoserts, M.H. and B.R. MacRoserts. 2004a. West Gulf Coastal Plain prairies:a first approximation at 
a synthesis. Third Eastern Native Grass Symposium, Chapel Hill, North Carolina: 5-21. 

MacRoetrts, M.H.and B.R.MacRoserts. 2004b, About 450 miles up the Red River in a large prairie: Peter 
Custis’ surviving botanical information from the Red River expedition of 1806. Sida 21:963-975. 

MacRoserts, M.H. and B. R. MacRoserts. 2005. Reference conditions of the Red River floodplain and 
upland, Caddo Parish, Louisiana. Sida 21:1793-1806. 

MacRoserts, M.H. and B.R. MacRoserrs (in prep.). Vascular flora and community structure of the C. 
Bickham Dickson Park/Red River Education and Research Park, Caddo Parish, Louisiana, Red River 
floodplain. 

Marsico, T.D. 2005. The vascular flora of Montgomery County, Arkansas. Sida 21: 2389-2423. 

McBryoe, J.B. 1933. The vegetation and habitat factors of the Carrizo sands. Ecol. Monogr. 3:247-297. 

Mclnnis, N.C., LLM. Smith, and A.B. Pittman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisi- 
ana. Phytologia 75:159-162. 

McKinney, LE. 1992. A taxonomic revision of the acaulescent blue violets (Viola) of North America. 
Sida Bot. Misc. 7:1-60. 

McKinney, L.E. and N.H. Russet. 2002. Violaceae of the southeastern United States. Castanea 67: 
369-379. 

McLaucutin, S.P.(in press). Tundra to Tropics: The floristic plant geography of North America. Sida, Bot 
Misc. 

Nesom, G.L.and L.E. Brown. 1998. Annotated checklist of the vascular plants of Walker, Montg y, 
and San Jacinto counties, east Texas. Phytologia 84: 107-153. 

Nesom, G.L. 2004a. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United 
States and comments on similar species. Sida 21:717-741. 

Nesom, G.L. 2004b. New distribution records for Gamochaeta (Asterac phali in the United 
States. Sida 21:1175-1185. 

Neyianp, R. 2002. An update to the vascular flora of Calcasieu Parish, Louisiana. Sida 20:431-433. 

Oversy, RE. 1974. A preliminary survey of the vascular flora of Caddo Parish, Louisiana. M.S. Thesis, 
Northeast Louisiana University, Monroe. 

Reio, C.and P. Fauckner (in press). Loeflingia squarrosa (Caryophyllaceae): new to Louisiana. Phytologia 

SKFEN, J.N., P.D. Doerr, and D.H. VAN Lear. 1993. Oak-hickory-pine forests. In: WH. Martin, S.G. Boyce, and 
A.C. Echtermacht, eds. Biodiversity of the southeastern United States. John Wiley, New York. Pp. 
1-33 


MACROBERTS AND MACROBERTS CADDO PARISH, LOUISIANA 1219 


Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Privately published. 
Fayetteville, Arkansas. 

Situ, E.B. 1994. Keys to the flora of Arkansas. Univ. Arkansas Press, Fayetteville. 

Teacue, J.and T. Wenor 1994. Caddo and Bossier parishes, Louisiana, natural areas survey. The Nature 
Conservancy, Baton Rouge. 

THomas, R.D.and C.M. Aiten. 1993-1998. Atlas of the vascular flora of Louisiana.Louisiana Department 
of Wildlife and Fisheries, Baton Rogue. 

THomas, R.D.,L.R. RavMonp, B.R. MacRoserts, and M.H.MacRoserts (in press). Documented and annotated 
checklist of the vascular flora of the Walter B. Jacobs Memorial Nature Park, Caddo Parish, Louisi- 
ana. Proc. Louisiana Acad. Sci. 

THorne, RF. 1993.Phytogeography. In: Flora of North America Editorial Committee, eds. Fl. North Amer. 
1:132-153.Oxford Univ. Press, New York and Oxford. 

Taiska, FJ. 1984. Role of wood debris in modifying channel geomorphology and riparian areas of a 
large lowland river under pristine conditions: a historical case study. Verh. Internat. Verein. Limnol. 
22:1876-1892. 

Turner, B.L., H.NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 
24:1-888. 


1220 BRIT.ORG/SIDA 22(2) 
Book REVIEW 
Doucias J. Kennet and Bruck WINTERHALDER. 20060. Behavioral Ecology and the Transition 
to Agriculture. (ISBN 0-520-24647-0, hbk.). University of California Press, Califor- 
nia/Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. 
(Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $60.00, 407 pp., 3 b/ 
w photos, 31 line illustrations, 22 maps, 30 tables, 7" x 10". 


Human behavioral ecology (as in the Foreword) focuses on how and why people have chosen, in social and cul- 
most funda- 


“This volume examines “one ol the m 


tural contexts, to exp dloit partic ular resources. From the Preface: 
mental economic shifts in human history—the evolutionary transition from foraging to farming through pro- 

he nenap tel authors use a variety of 
cen- 


cesses ? pen ane mee sscuiiat iaetcac ane. the emergence A apiculate. T 


spersed cz 

tral ee betas citeuncee. risk minimization, antl costly signaling shieae y. Their euanibirens are novel in 

presenting regionally comprehensive case studies that address the transition to agriculture from a consistent 

conceptual framework informed by neo-Darwinian theory.” A collection of 14 technical but readable papers.— 
Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S 


Book Notice 
W. SHENG YI, PeTeR H. Raven, and H. Deyuan (editorial co-chairs). 2006. The Flora of China, 
Volume 22, Poaceae. (ISBN 1-930723-50-4, hbk.). Missouri Botanical Garden Press, 
P.O. Box 299, Saint Louis, MO 63166-0299, U.S.A. (Orders: 877-271-1930, http:// 
www.mbgpress.org) $140.00, 752 pp.,9" x 111/2" 
le family, the Poaceae, ene 


-volume work. : umerades sing ) 


] 809 < 


eas 22 of ne E ne of China isthe l2th of a 25 
tribes, 226 genera, and 1795 species, among 

to China.” FE xcept for the protracted list of Asian Bambuseae genera (34), me sity in genera berieene a 

and North America is remarkable. The Chinese grass flora also is close in size to the North American one (nor 

of Mexico), perhaps even 20% greater in number of species. The first of two aes of oe FNA treatment was 

Many recent cl ts of grass genera are incor- 

the FNA treatments.—Guy 


published in 2003; it will be published in early 2007. 
porated in the Chinese flora, presumably making it parallel and peers sith 
Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. 


= 
rt 


SIDA 22(2): 1220. 2006 


CAREX STIPATA (CYPERACEAE), CHAMAESYCE GEYERI 
(EUPHORBIACEAE), EURY TAENIA TEXANA (APIACEAE), 
PEDIOMELUM ESCULENTUM (FABACEAE), AND TALINUM 
CALYCINUM (PORTULACACEAE) DELETED 
FROM THE LOUISIANA FLORA 


Barbara R. MacRoberts and Michael H. MacRoberts 


Bog Research, 740 Columbia, Shreveport, Louisiana 71104, U.S.A. 
Herbarium, Museum of Life Sciences, Louisiana State University-Shreveport 


Shreveport, Louisiana 71115, U.S.A. 


anc 
Red River Watershed Management Institute, Louisiana State University-Shreveport 
Shreveport, Louisiana 71115, U.S.A. 


In the course of developing a vouchered checklist of the vascular flora of Caddo Parish, 
Louisiana (MacRoberts & MacRoberts in prep.), we discovered six species reported for 
Caddo Parish for which we could find vouchers neither for the Parish nor the State. We 
already have reported on one of these: Agrimonia gryposepala Wallr. (MacRoberts & 
MacRoberts 2005). Here we report on five others. Until vouchers are found, these species 
should be deleted from the Louisiana flora. 

Carex stipata.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), 
Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005) include Carex 
stipata Muhl. ex Willd. in the Louisiana flora based on a specimen collected by D.T. 
MacRoberts in Caddo Parish (MacRoberts 1349 LSUS). We examined the specimen and 
found it to be a misidentified Carex vulpinoidea Michx. (Phil Hyatt also annotated it to C. 
vulpinoidea). The reported distribution of C. stipata is confused. It is not reported for Loui- 
siana, Mississippi, or Arkansas in the Flora of North America but it is reported for these 
states by Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005). Turner 
et al. (2003) do not report it for Texas. 

Chamaesyce geyeri.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), 
Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005) include Chamaesyce 
geyeri (Engelm.) Small in the Louisiana flora ae on several specimens collected by 
D.T. MacRoberts in Caddo Parish (MacRoberts 1791, 1940 LSUS, 1951 LSU). We examined 
these specimens and found them to be misidentified Chamaesyce cordifolia (EIL.) Small. 
Chamaesyce geyeri is a central plains species that apparently does not occur nearer to 
Louisiana than central Texas and central Oklahoma (Turner et al. 2003, Oklahoma Vas- 
cular Plant Database 2005). 

Eurytaenia texana.—Thieret (1971), MacRoberts (1979, 1984, 1989), Thomas and Allen 
(1993-1998), Kartesz and Meacham (1999), LSU (2005), NatureServe (2005), and US.D.A. 
(2005) include Eurytaenia texana Torr. & A. Gray in the Louisiana flora on the basis of 
many collections from Caddo, Winn, Webster, and DeSoto parishes (e.g., [hieret 32309 
LAE Thomas 45467, 65016, 70521, 99542, 99703 NLU, Thompson 485 LTU, Dixon 1342 NLU). 
We examined these specimens and others labeled Eurytaenia texana from Louisiana in 
BRIT, NLU, LSU, LTU, and LAF and found all of them to be misidentified Polytaenia 


SIDA 22(2): ): 1221-1223. 2006. 


1222 BRIT.ORG/SIDA 22(2) 


nuttallii DC. The distribution of Eurytaenia texana is central Texas, the Texas coastal 
prairie region, and western Oklahoma (Mathias & Constance 1961, Taylor & Taylor 1989, 
Hatch et al. 1990, Turner et al. 2003, Oklahoma Vascular Plant Database 2005, TAMU 
2005). It is, however, reported from Jefferson Co., Texas, on the Louisiana Gulf border and 
thus could possibly be present in the Louisiana coastal prairie region (Mathias & Con- 
stance 1961, Turner et al. 2003). A reported specimen of Eurytaenia texana from Arkan- 
sas housed at NLU turned out to be a misidentified Thaspium barbinode (Michx.) Nutt. 
(Eric Sundell, pers. comm.). 

Pediomelum esculentum.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993- 
1998), Kartesz and Meacham (1999) and U.S.D.A (2005) include Pediomelum esculentum 
(Pursh) Rydb. in the Louisiana flora based on a specimen collected by D.T. MacRoberts 
from Caddo Parish (MacRoberts 1170 LSUS). We examined the specimen and found it to 
be a misidentified Pediomelum hypogaeum (Nutt. ex Torr. & A. Gray) Rydb. var. subulatum 
(Bush) J. Grimes. Pediomelum esculentum apparently does not occur nearer to Louisiana 
than northern Arkansas and central Oklahoma (Oklahoma Vascular Plant Database 2005, 
US.D.A. 2005). 

Talinum calycinum.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), 
Kartesz and Meacham (1999), NatureServe (2005) and US.D.A (2005) include Talinum 
calycinum Engelm. in the Louisiana flora based on specimens collected by Thomas in 
Caddo and Vernon parishes (Thomas 34513, 105079 NLU). We examined all Louisiana T. 
calycinum specimens (MacRoberts & MacRoberts 1997) and found them to be 
misidentified I’ parviflorum Nutt. Talinum calycinum apparenty does not occur nearer 
to Louisiana than central Texas, Oklahoma, and central Arkansas (Turner et al. 2003, 
Oklahoma Vascular Plant Database 2005, U.S.D.A. 2005). 


ACKNOWLEDGMENTS 


Diane Ferguson (LSU), Kerry Heafner (NLU), Garrie Landry (LAF), Amanda Neill (BRIT), 

Eric Sundell (UAM), John Thieret (KNK), Milan Vavrick (LTU), and Gary Hanson and 

Patricia Peyton, Red River Watershed Management Institute, aided with this work. R. Dale 

Thomas kindly reviewed the paper. 

REFERENCES 

Harcn, S.L, KN. GANDHI, and L.E. Brown, 1990. Checklist of the vascular plants of Texas. Texas Agricul- 
tural Experiment Station, Texas A&M University, College Station. 

Karresz, J.T. and CA. Meacuam. 1999. Synthesis of North American flora. Version 1.0 North Carolina 
Botanical Garden, Chapel Hill. 

LSU 2005. www.herbarium.|su.edu 

MacRoserts, B.R.and M.H. MacRosrers. 2005. Agrimonia gryposepala (Rosaceae) deleted from the Loui- 
siana flora. Phytologia 87:129-131. 

MacRoserts, B.R. and M.H. MacRoseets. in prep. Vouchered checklist of the vascular flora of Caddo 
Parish, Louisiana, with notes on regional phytogeography and ecology. 

MacRoseets, D.T. 1979. Checklist of the plants of Caddo Parish, Louisiana. Bull. Mus. Life Sci. Louisiana 
State Univ. 1:1—54. 

MacRoserts, D.T. 1984. The vascular plants of Louisiana. An annotated checklist and bibliography of 
the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. Louisiana 
State Univ. 6:1-165. 


FLORA 1223 


MacRoseats, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus 
Life Sci. Louisiana State Univ. 8:257-536. 

MacRoserts, M.H.and B.R. MacRoserts. 1997. Talinum rugospermum Holz.,new to Louisiana with notes 
on terete-leaved Talinum in Louisiana. Phytologia 82:86-93. 

Martuias, M.E. and L. Constance.1961. Umbelliferae. Flora of Texas. Vol. 3. Part 5. Texas Research Founda- 
tion, Renner. 

NatureServe. 2005. www.natureserve.org 

OKLAHOMA VASCULAR PLANT Database. 2005. http://g botanical/ 

TAMU 2005. www.csdl.tamu edu/FLORA/tracy2/main| hel 

Tayior, R.J.and C.E.S. Tayior. 1989. An annotated list of the ferns, fern allies, gymnosperms and flower- 
ing plants of Oklahoma. Southeastern Oklahoma State University, Durant. 

Tuieret, JW. 1971. Additions to the Louisiana flora. Castanea 36:219-222. 

THomas, R.D.and C.M. Atten. 1993-1998. Atlas of the vascular flora of Louisiana. Louisiana Department 
of Wildlife and Fisheries, Baton Rouge. 

Turner, B.L., N. NicHots, G. Denny, and O. Doran. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 
24:1-888. 

U.S.D.A. 2004. http://plants.usda.gov 


1224 BRIT.ORG/SIDA 22(2) 


Book NOTICES 


Yonc No Ler. 2006. New Flora of Korea (two volume set). ISBN 89-09-11801-6, vol. 1: ISBN 
89-09-11802-4, vol. 2, hbk.). Kyo-Hak Publishing Co., Ltd., 105-67, Gongdeok-dong, 
Mapo-gu, Seoul, Korea. (Orders: 02-7075-155-156, 02-7075-160 Fax, chief @k yohak.co.kr, 
www:kyohak.co.kr) $345.00 (postage (surface) included to U.S.A.), 976 pp., vol. 1; 888 
pp., vol. 2, abundant color Maia Ox Ly 2 


So many beautiful and detailed color photos (1 estimate 80+% of the species with a photo: match photo number 


yer)... but Lcan read the scientifien names, that’s all. Ferns, grasses, sedges, trees, daisies, and all 


with species num 
the rest—all illustrated in two big volumes (boxed set). It’s not cheap but what a value for botanists. From the 
‘flyer’ 4157 taxa of native and naturalized plants of Korea; almost all plant pictures have been taken by the au- 
thor, Vol. | with 976 pages, Vol. I] with 888; scientific names, proper Korean names, English names, Chinese names 


and Japanese names have been used; habitats, distributions, flowering seasons, and chromosome numbers have 


been recorded; new species and rearranged species names have been arranged in an appendix pinay an in- 
dex of new taxa and new combinations by the author in various literature from 1993 to early 2006, joined by 
many new combinations proposed here de novo). Since synonyms are given for very few of the scientific names, 


the index to scientific names functions as a checklist for the whole flora —Guy Nesom, Botanical Research Insti- 
tute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. 


Davin Baxter (Photographs by Laurence Parent). 2002. Nature of the Forest: Temple-Inland’s 
Timberlands in the Twenty-first Century. (ISBN 009718675-0-X, hbk.). Temple-In- 
land Inc. No other information provided. 


It's not easy to be enthusiastic about books about timber companies, published by timber companies, because so 
much private forest land has been turned to monoculture, the diversity of all plants reduced literally to a few 
species. Practices of the USDA Forest Service commonly give the same result over large portions of public land. 
This book obviously is intended to put the best face possible on the Temple-Inland timber company—nicely written 
is surely one of the white Gwhiter?) hats ces . ba: lates in Hmber 


and full of beautiful pictures. But T 
management, as it apparently has set aside an unusual amount of land for preservation 
The book notes that “Temple maintains about a quarter of its land as natural forest;” .. some seventeen percent af 
its land is in SMZs [streamside ne ment zones|—340,000 acres. Plans are to level olf at eighteen to twenty 
percent of the company’s total acreage.” Italso maintains a number of “special places” conserved within its hold- 


ings in east Texas, Alabama, and Georgia, and in 2006 the Native Plant Society of Texas recognized Temple-In- 


ynatur 


Qa 


with an award for its contribution to native plant habitat preservation in East Texas. 
As noted in the book, “Temple-Inland is in the tree puSnieSS Bu une greater Temple-Inland company has 


lanc 


expertise in real estate ove opment As such, Temple is company land for its ‘highest 


and best use.’ Temple already owns the land, so would not be out the cost ay is cai — might ~~ at oni 
ly 


types of development.” The ultimate fate of TI holdings and all timber lands has recent 

unpredictable—Boise-Cascade sold all of its 2.3 million acres of forestland in 2004, and in 2005 ineeroationel 
Paper announced the sale of its 6.8 million acres of North American timberland, including many conservation 
areas (see Nature Conservancy, Autumn 2006). Only a small fraction of this land has been moved into some kind 


of conservation status. At least commercial timber land, even with its greatly reduced species diversity, can be 
coaxed back into a semi-natural condition, but other kinds of development (housing or commercial) are more 
nearly irrevocable.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, 
U.S.A 


SIDA 22(2): 1224. 2006 


NEW VASCULAR PLANT RECORDS FOR NEW MEXICO 


Ronald L. Hartman Brian Reif 
Rocky Mountain Herbarium 53 Chof Trail 
Department of Botany, Dept. 3165 Flagstaff, Arizona 86001, U.S.A. 
ae versity of pa ing Brianreif_2000@yahoo.com 
000 E. University Ave. 
Laramie ‘ee 82071, U.S.A. 
Rhartman@uwyo.edu 
B.E. Nelson Brian Jacobs 
Rocky Mountain Herbarium ource Management 
mites ae Dept. 3165 na National Monument 
University of Wyoming National Park Service 
1000 - eee ity Ave. HCR-1, Box 1, Suite 15 
Laramie Wyoming 82071, U.S.A. Los Alamos, New Mexico 87544, U.S.A. 
Bnelsonn@uwyo.edu Brian_jacobs@nps.gov 
ABSTRACT 


Additions to the vascular flora of New Mexico are reported herein: Achnatherum nelsonii var. sada ie 


glandulosa, Cardamine cordifolia var. incana, Carex deweyana var. deweyana, Carex rosea, Cynos 
Draba cate Equisetum Xnelsonii, Erigeron nivalis, Geum triflorum var. triflorum, ene hallii judians 
nigra, Lactuca biennis, Lepidium ramosissimum var. bourgeauanum, Neoparrya lithophila, Penstemon glaber var. 

alpinus, Piptatheru gens, Potentilla fissa, Prunus persica, Ranunculus alismifolius var. montanus, Ranuncu- 


lus repens, Renibod yes Rudbec kia laciniata var. laciniata, and Syringa vulgaris. 


RESUMEN 


\chnath lsonii var. nelsonii, Bet ula 


-] 


Se aportan las adiciones siguientes a la flora vascular de Nuevo México: 
glandulosa, Cardamine cordifolia var. incana, Carex deweyana var. dewe yana, Carex rosea, Cyn 
Draba grayana, Equisetum Xnelsonii, Erigeron nivalis, Geum t triflorum var. triflorum, eee inal it; ue 
nigra, Lactuca biennis, Lepidium ramosissimum var. bourgeauanum, Neoparrya lithophila, Penstemon glaber var. 
alpinus, Piptatherum pungens, Potentilla fissa, Prunus persica, Ranunculus alismifolius var. montanus, Ranuncu- 
lus repens, Rorippa sylvestris, Rudbeckia laciniata var. laciniata, y Syringa vulgaris 


Past and recent inventories on the flora of north central New Mexico have resulted in 24 
native or naturalized taxa new to the state or verification of reports. Much of this work is 
part of an ongoing inventory of the Rocky Mountains by the students and staff of the 
Rocky Mountain Herbarium (http://www.rmh.uwyo.edu). Thus far more than 44 inten- 
sive, large-scale projects have been completed. More specifically, these records are the re- 
sult of the following inventories: Philmont National Scout Ranch (Hartman 1973), 
Bandelier National Monument (Jacobs 1989), Valles Caldera National Preserve (Hartman 
& Nelson 2005), and the Santa Fe National Forest (Reif @ Hartman 2005; Reif 2006). An 
additional study, Carson National Forest and adjoining Bureau of Management lands, is 
in its second year (Larson, Hartman, and Nelson, unpub.). All specimens are deposited at 
the Rocky Mountain Herbarium (RM) or at the herbarium of Bandelier National Monu- 
ment (here designated BAND). Determination that a taxon is new to New Mexico is based 
on Allred (2005). In several instances a taxon included in this report has been attributed 
to New Mexico, but without documentation. 


SIDA 22(2): 1225- 1233. 2006 


1226 BRIT.ORG/SIDA 22(2) 


APIACEAE 

Neoparrya lithophila Mathias—The type locality for this species was attributed to New 
Mexico (“on rocks, Huefano[Huerfano] Mts, New Mexico, Sept., 1867, CC. Parry 83”) but is 
now considered to be from Huerfano Co., Colorado (Weber 1958, Theobald et al. 1964). 
Hartman (in O'Kane et al. 1988) reported it from several sites as far as 51 mi to the south- 
west and 70-80 mi to the west of the type locality, previously the only known site for the 
species. Based on this report and subsequent inventory in San Isabel and Rio Grande Na- 
tional Forests and vicinity, it is now documented from more than 30 localities largely 
along the eastern margin of the San Juan Volcanic Area. Most recently, collections (RLH) 
have been made in the South Pifion Hills in the extreme southern San Luis Valley, imme- 
diately to the north of the locality reported below. Thus a taxon originally attributed 
erroneously to New Mexico, then considered endemic to Colorado, indeed occurs in New 
Mexico but only by one-half mile. 


— 


Voucher specimen: Taos Co.: Carson National Forest and Vicinity: southern extension se Pinon Hills into New 
Mexico, N36.9915 W105.7935, 7900 ft elev, rocky outcrop, 29 Jul 2005, Hartman 8159 { 


ASTERACEAE 
Erigeron nivalis Nutt.—The following specimen was included in Allred (2005) as E. acris 
Var. asteroides (Anderz. CX Besser) DC. due to a misidentification (Hartman 2003). In the 


United States, E. nivalis ranges from the Rocky Mountains to the West Coast excluding 
Arizona and Nevada. It was attributed to New Mexico by Nesom (2006) based on the 
Ludwig and Smith and the Reif collections cited below. 


Voucher specimens: Sandoval Co.: Baca Land and Cattle Company, Redondo Peak, N35.87 W 106.66, 10925 ft 
elev, talus slope, 1-9 Jul 1975, J. Ludwig and S. Smith 1368 (NMC): Valles Caldera National Preserve, Jemez Moun- 
tains: upper slopes on NE side of Redondo Peak, N35.8806 W106.5484, 10160-10360 ft elev, roadside along volca- 
nic outcrop, open slopes, and forested areas, 15 Jul 2002, Reif 2403 (NMC, RM); trip from 7 roads to “Redondito 
peak” and down the La Jara Creek to headquarters: 0.5-0.7 air mi ENE of 7 roads, N35.8923 W106,5536 to N35.8908 
W 106.5480, 9730- i ft elev, spruce-tir forest and adjacent felsenmeer and g grown over 2-tracks, 18 Jul 2003, 
Hartman 77974 (RM 

Lactuca biennis a Fernald—USDA, NRCS (20006) maps this taxon for New Mexico 
but without voucher data and it is not included in Allred’s checklist (2005). Thus docu- 
mentation is provided below. 

Voucher specimens: Colfax Co.: Philmont Scout Ranch: Middle Fork Cimarroncito Creek, 1 mi W of Hunting 
Lodge, N36.4844 W105.0801, 8200 ft elev. exposed waste run along road, 7 Aug 1968, Hartman 2626 (RM). San- 
doval Co.: Rito de los Frijoles, N35.7793 W106.2719, Aug 1910, Robbins 8210 (RM). Bandelier National Monument: 
Frijoles Canyon, ca. 2 mi below Upper Crossing, N35.8044 W106.3301, 6500 ft elev, moist canyon bottom ans 
stream, 24 Jul 1988, B. Jacobs and E. Jacobs 4353 (BAND); 20 Jul 1941, Clark 9935 (BAND). San uel C 

N35.8050 W105.4407, 8000 ft elev, Aug 1902, Cockerell 11 (RM). Santa Fe National Forest and Vicinity: Sangre ae 
Cristo Mountains: Maestas Creek on NM hwy 391, 4.5 air mi we ot Rack ada, N35.8568 W105. ee 8500- 8680 


ft elev, riparian and moist banks of creek, 15 Aug 2002, Reif 4 cos Wilderness Area, Holy Ghost Creek, 
3 air mi NW of Terrero, N35.7856 W105.7048, 8100-8360 ft elev, eee e meadow in canyon bottom, 28 Jul 
2002, Reif 2768 (RM) oie Canyon on NM hwy 123 branching W of the Pecos River, 6 air mi NNW of Pecos, 
N35.6712 W105.7412, 7760 ft elev, wet to dry meadow and riparian with canyon constricting to the W, 6 Aug 
2002, Reif 3392 (RM) 


Rudbeckia laciniata L. var. laciniata— Variety ampla(A. Nelson) Crongq. is not distinguished 
[rom the typical variety in Allred (2005). The former is the common variety throughout 
the Rocky Mountains with var. laciniata entering the region from the Great Plains. USDA, 
NRCS (2006) indicates its occurrence in New Mexico but without documentation. Thus 


HARTMAN ET Al 1227 


var. laciniata is vouchered below for the state and the reader is referred to Urbatsch and 
Cox (2006) for the discrimination of the two taxa. 
Voucher specimens: Colfax Co.: NM hwy 72, ll rd mi E i -25 (milepost 11), N36.9219 W104.2783, 8200 ft elev, 
roadside through wooded area, 16 Aug 1998, Hartm RM). San Miguel Co.: Santa Fe National Forest and 
Vicinity: Sangre de Cristo Mountains: NM hwy 263 atone Gallinas Creek, 16 air mi NW of Las Vegas, N35.7083 
W105.4505, 7750 ft elev, canyon bottom roadside, riparian, and bluffs of volcanic rock in mixed conifer forest, 13 
Aug 2002, Reif 3820 (RM). Santa Fe Co.: Santa Fe National Forest and Vicinity: Sangre de Cristo Mountains: Rio 
Frijoles Trail 154 just E of Santo Domingo de Cundiyo Land Grant, 8 air mi SE of Chimayo, N35.9380 W105.8733, 
00 ft elev, riparian of river, and meadows, in canyon bottom with mixed conifer slopes, 8 Aug 2002, Reif 3421 
(RM) 


BETULACEAE 

Betula glandulosa Michx.—Furlow (1997) shows this taxon extending down the Rocky 
Mountains to south-central Colorado. Its presence on the Valles Caldera National Pre- 
serve was first noted by Allen (2004) who found numerous stunted birch plants growing 
on Alamo Bog. This site represents a disjunction of 100 miles or more from the nearest 
known location in Colorado (http://cumuseum.colorado.edu). Heavy browsing by elk 
appears to be jeopardizing the population. 

Voucher specimens: Sandoval Co.: Valles Caldera National Preserve: Jemez Mountains: Alamo Canyon, 1-15 mi 
ae N35.9182 W106.6032, 8461ft elev, over 43 healthy but cropped shrubs to 2' tall along 1/4 mi of shore- 

south side, 2 Jun 2003, Hartman 77223 (RM); 8 Jun 2001, Jacobs 14 (BAND). 


BRASSICACEAE 

Cardamine cordifolia A. Gray var. incana A. Gray ex ME. Jones—This markedly pubes- 
cent variety is quite distinctive when compared to the essentially glabrous typical one 
(Rollins 1993; Welsh et al. 2003). Holmgren (2005) does not recognize var. incana but notes 
that the epithet is available for the pubescent form. Previously it was known from Colo- 
rado and Utah. 

Voucher specimen: Rio Arriba Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, upper Cananones 
Creek and W slope, 11. 3 air mi S of Youngsville, N36.0304 W106.5422 to N36.0342 W106.5271, 9300-10000 ft 
elev, springs on trail in | Pp kbark fir forest with mature aspen, 25 Jun 2002, Reif 1197 (RM 


Draba grayana (Rydb.) CL. Hitchc—This taxon has been considered endemic to Colo- 

rado, occurring in its northern and central portions (Rollins 1993). With its discovery on 

Little Costilla Peak, it is now documented from New Mexico. 

Voucl olfax Co.: Sangre de Cristo M , Carson National Forest- Valle Vidal Unit, Little Costilla 
Peak, Beiec N36.8260 W105.2234 and N36.8336 W105.2228, 12,231-12,584 ft elev, alpine slopes, 16 Aug 2002, 

Hartman 76640 (RM). Determined by Ihsan Al-Shehbaz (MO) 


Lepidium ramosissimum A. Nelson var. bourgeauanum (Thell.) Rollins—In the western 
United States, this variety ranges from Montana south to Colorado and west, including 
Arizona (USDA, NRCS 2006). According to Rollins 993) “the varieties here recognized 
are weak at best. The consistently glabrous siliques of var. bourgeauanumare not matched 
by consistency in var. ramosissimum, where the siliques are mostly sparsely pubescent, 
but on occasion both pubescent-fruited and glabrous-fruited plants are found in the same 
population. The chromosome difference of 2n = 32 for var. bourgeauanum and 2n = 64 for 
var. ramosissimum. .. helps to bolster the treatment where the two varieties are recog- 
nized... .” Holmgren (2005) accepts the two varieties without discussion. We maintain 
the two taxa pending further study. 


1228 BRIT.ORG/SIDA 22(2 


Voucher specimens: Sandoval Co.: Valles Caldera National Preserve, Jemez Mountains, Jemez Falls Road to turn 
iwack to southwest corner of Preserve, N35.8277 W100.6170, 8000-8164 [t elev, ponde- 
rosa pine forest with rock outcrops and swales, 30 Jul 2001, ase 73133 (RM); just S of Sulphur Point, N35.9116 
W 106.6213, rocky slopes in mixed conifers, 8981 [t elev, 17 Sep 2001, Hartman 74216 (RM); Alamo Canyon, south 
side over a mile stretch to near lower end, N35.9117 W106. 5002 to N35.9109 W106.5927, 8650-8960 ft elev, scat- 
tered at edge of wet area over lower half of boggy portion, 19 Jul 2003, Hartman 78125 with Coop (RM); Puerta de 
Trasquilar NW up end of Trasquilar, N35.9398 W106.4960 to N35.9423 W106.5002, 8925-9360 ft elev. south ex- 
posure with oak and aspen, 24 Jul 2003, Hartman 78203. Puerta de Trasquilar NW up end of Trasquilar, N35.9398 
W106.4960 to N35.9423 W106.5002, 8925-9360 ft elev, south exposure with oak and aspen, 24 Jul 2003, Hart- 
man 78234,(RM); Valle Jaramillo, near “B” and “M” road intersection, N 35.9184 W106.4981 to N35.9133 W106.4912, 
8705-8720 ft elev. grassy meadow with creek and mud areas on slope with upwelling of water, 24 Jul 2003, 
Hartman 78258 (RM); Valle Toledo, crossing of Pipeline Road and San Antonio Creek to 0.4 air mito NE then S$ to 
0.3 air mi ae N35.9602 W106.4827 to N35.9613 W106.4736, 8610-8639 [t elev, wetlands along creek to 


1] 


~ 


south, two track and bus 


— 


Qo. 


slopes to N w vith barren hau and adjacent “bouncy” ground, 10 Aug 2003, Hartman 


78348 (RM); on Fork ae River ae segment in Preserve ca. 1 air mi SW of El Cajete (center of caldera), 
N35.8277 W106.5751 to N35.8277 W106.5780, on 7990 ft elev. along river with blue spruce and white fir on 
slopes, willow and birch scattered on bank, Ll Aug 2003, Hartman 78425(RM); Banco Bonito, 0.8-0.9 air mi NE of 
southwest corner of Preserve, N35.8385 W106.6136 to N35.8367 W106.6078, 8250-8332 ft elev., roadside through 
ponderosa pine, 12 Aug 2003, Hartman 78449 (RM); Pajarito Mountain, ca. | air mi WNW of summit, N35.8897 
W106.4172 to N35.8897 W106.4173, 8970-9050 ft elev. lower grassy slopes to scattered ponderosa pine, 21 Aug 
2003, Hartman 78802 (RM); off of G Road just above F Road, ca. Lair mi N of Cerro Santa Rosa, just N of Valle San 
Antonio, N35.9743 W106.5050, 8560-8800 [t elev, logged ponderosa pine forest, 5 Aug 2003, Nelson 59659(RM): 
Jemez Mountains, from 3/4 mi E to N35.9775 W106.4827 in Rito de los Indios drainage, 8750-9000 ft elev, dis- 
turbed areas, 5 Aug 2003, Nelson 59700 (RM); from tributary of Rito de los Indios NW to summit of ridge, ca. 1 1/ 
2-1 3/4 air mi N of north summit of Cerros de Trasquilar, N35.9921 W106.5020, 9100-9740 ft elev, open, grassy 
slopes with scattered ponderosa pine, 5 Aug 2003, Nelson 59757 (RM); along G Road ca. 1 3/4 mi N of F Road, ca. 
2 1/2-2 3/4 air mi N of Cerro Santa Rosa, N35.9978 W106.5006, 9600-9800 . elev, meadow, 5 Aug 2003, Nelson 
59831(RM); on the ridge W of Valle Grande, ca. 11/4-1 3/4 air mi N of Ranch Headquarters, N 35.8806 W106.5180, 
8900-9600 ft elev, roadside through mixed conifer forest, 7 Aug 2003, Ne en 59942 (RM); ridge S and E of Valle 
Jaramillo, ca. 1 air mi NW of Cerro Pinon, N 35.8993 W106.5124, es ft Bie clearcut Douglas fir forest with 
mos stly | blue spruce white fir regeneration, LO Aug 2003, Nelson 60025 (RM): a g K Re yad at the northeast base 
of San Antonio Sane ain, N35.9449 W106.5925 to N35.9437 W106. 5089, 8000- 9100 ft elev, roz mae and adja- 
cent open spen forest, 1] Aug cose Nelson ep0ee (RM); along the northeast and north side of San 
Antonio Mountain to the west boundary, N35.94 to N35.9463 W106.6204, 900¢ ee along 
old logging road, mixed conifer/aspen forest, a ae 1 Aug 2003, Nelson 60118 (RM); on the east slope of 
Cerros del Abrigo above the Abrigo aa : 0. 3mi on either side, N35.9301 W106.4726, ee ft Bley, i 


0314(R 


old logging roa spen forest, and open areas,13 Aug 2003,) on 
south side of Cerros del Abies along the Ahaee Trail for 0.3 mi on either side, N35.9232 W106.4768, 9400 ft elev 
along old logging road, roadcut, and adjacent southwestern white pine/white fir/Douglas fir forest. 13 Aug 2003. 
Nelson 60243 (RM); Santa Fe Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Rio Frijoles 
Trail 154 just E of Santo ati de Cundiyo Land Grant, c air mi SE . Chimayo, N35.9380 W105.8733, 7000 ft 
elev, riparian of river, and meadows, in canyon bottom v r slopes, 8 Aug 2002, Reif 3448 (RM). 


Rorippa sylvestris (L.) Besser—Introduced throughout much of the United States, this 
mustard is said to occur in New Mexico: “reported questionably by Kartesz and Meacham 
(1999): awaits verification” Allred (2005). 

Voucher specimen: San Miguel Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Anton 
Chico Grant, 5 air mi ENE of Villanueva, along Pecos River, 35.2365 105.2558, 5500 ft elev, river bottom, mostly 
juniper with Russian olive and narrowleat cottonwood, 6 Aug 2004, Reif 10067 (RM). 


CYPERACEAE 

Carex deweyana Schwein. var. deweyana—Naczi (2002) lists but does not map this taxon 
flor New Mexico, although it is mapped for the state in USDA, NRCS (2006). Interesting]y, 
it isnot included in Allred (2005). Regardless, the first two sources do not contain voucher 
data. Consequently, our records are provided below. 


HARTMAN ET Al 1229 


Voucher specimens: Los Alamos Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, lower Guaje 

yon and Cabra Canyon, 3 air mi N of Los Alamos, N35.9310 W106.3432 to N35.9064 W106.2862, 7000- a ft 
elev, riparian drainage through mixed conifer forest, into a burn past the canyon, and then a ponderosa pine 
forest before joining the road, 28 Jun 2002, Reif 1475 (RM). Santa Fe Co.: Santa Fe National Forest and Vicinity, 
Sangre de Cristo Mountains, Pecos Wilderness, Rio Merlo tal 155 acincent to Rio Medio, 6 air mi due W of Truchas 
Peak, N35.9630 W105.7520, 8410 ft elev, rip 

or standing water, 23 Jun 2003, Reif 5806 (RM). 


> of lower canyon slopes, banks of river 


(=) 


Carex rosea Schkuhr ex Willd. According to Ball (2002), this taxon ranges from Wyo- 
ming, the Great Plains and Texas to the east coast of North America. 


Voucher specimens: Sandoval Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, Cochiti Canyon, 4.3 

air mi SSE of Rabbit Mountain, N35.7794 W106.4366 to N35.7645 W106.4266, 7200-8280 ft elev, canyon bottom 

and riparian, 23 Jun 2003, Reif 1116 (RM). San Miguel Co.: Santa Fe National Forest and Sana Sangre de Cristo 

ee. Sebadilla Creek S of Flying Cloud Ranch, 3 air mi SSW of Barillas Peak, u ek 1/2 air mi, N35.5320 
7740-7880 ft elev, narrow riparian corridor in canyon bottom, canyon al by mixed are 

forest, sean bluffs and hee constrictions, 9 Jun 2003, Reif 5070 (RM); El Porvenir Canyon on Trail 247, 

mi W of San Ignacio, N35.770 W105.448, 8200-8450 ft elev, canyon bottom of mostly spruce, and moist ie 


slope of mixed conifers along am granite bluffs constricting canyon, 6 Jul 2003, Reif 6183 (RM). 


EQUISETACEAE 

Equisetum Xnelsonii (A.A. Eaton) J.H. Schaffn.—This plant represents a spontaneous 
hybrid between E. laevigatum and E. variegatum. The former putative parent is frequent 
in the Jemez Mountains; E. variegatum is not documented from New Mexico but has been 
collected as far south as Montezuma County, Colorado, some 130 mi to the northwest. In 
the Rocky Mountains, the hybrid is known from Montana, Wyoming, Utah, and Colo- 
rado (USDA, NRCS 2006) 

Voucher specimen: Sandoval Co.: Bandelier National Monument, White Rock Canyon at mouth of Alamo Can- 
yon along banks of Rio Grande, N35.7146 W106.2907, 5300 ft elev, 23 Aug 1975, Foxx s.n.(B ) 


JUGLANDACEAE 
Juglans nigra L—Black walnut, a native of the eastern United States, has been cultivated 
in areas of the western United States. As described below, it is naturalized in NM. 


J 


Voucher specimens: Sandoval Co.: Bandelier National Monument, vicinity of P 5 in Canyon de los 
Frijoles, along Rito de los Frijoles near and below Tyuonyi Ruin, N35.7839 W106.2772, 6, 100 ft elev. escaped third 
peneration seceine meattcree in colconnond bosqt ue alae stream, os Sep ee Jacobs 5083 (BAND), second gen- 

historic Frey Abbott orchard, 4 May 2003, Jacobs 5084 (BAND), 


OLEACEAE 

Syringa vulgaris |.—This introduced lilac occurs over much of eastern United States and 
scattered states in the West. The latter includes Kansas, Colorado, and Utah (USDA, NRCS 
2006). This collection appears to be an adventive representing the first New Mexico record. 


Voucher ae ae Santa Fe Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Tesuque 

n Winsor Trail 254, 3 air mi SE of Tesuque, N35.745 W105.888 SW to N35.743 W105.896, 7320-7370 ft 
ae narrow canyon bottom and riparian with mature ponderosa pine, horsetail, cotton wood, alder, and willow, 
granitic soils, 12 Jun 2003, Reif 5252 (RM). 


POACEAE 

Achnatherum nelsonii Scribn. var. nelsonii— According to Barkworth et al. (1979), this grass 
ranges from southern Montana and central Idaho south through Nevada and Utah. In 
Utah it has been documented in the southeast corner, immediately adjacent to New 
Mexico. In addition, USDA, NRCS (2006) maps it for Colorado and Arizona. Verified by 
Mary E. Barkworth. 


1230 BRIT.ORG/SIDA 22(2) 


Voucher specimens: Rio Arriba Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, vicinity of Presa 
ing on southern edge of Chama River Canyon Wilderness, 8 air mi NE of Gallina, N36.3120 W106.7340, 8340 

elev, a forest of mature ponderosa pine, with thickets of Gambels oak and ao open areas below, 19 Jun 2003, 
- if 5617 (RM). Sandoval Co.: Sierra Nacimiento, Joaquin Canyon, ca. 2 3/4-3 1/4 air mi W of Rio Guadalupe, 
N35.79 x W106.8 oh coe 8050 ft elev, open area along logging road and adjacent mixed conifer forest, 20 Jul 
2004, b 621 


Cynosurus echinatus L. This grass has been introduced in much of eastern United States, 
the West Coast, and Montana (USDA, NRCS 2006). This represents the first report of its 
establishment in New Mexico. 

Voucher specimen: Sandoval Co.: Bandelier National Monument, Capulin Canyon, ca. 0.25 mi S of Base Camp, 
near where trail to Stone Lions joins canyon trail, N35.7541 W106.3262, 6100 ft elev, dense stand along stream, 13 
Jun 1988, B. Jacobs 4167 (BAND). 

Piptatherum pungens (Torr. ex Spreng.) Dorn—This species ranges across western Canada 
and south in the Black Hills of Wyoming and South Dakota to the central cordillera of 
Colorado (Kartesz and Meacham 1999, USDA, NRCS 2006). In Colorado it is documented 
from Douglas, Grand, and Saguache cos. (http://cumuseum.colorado.edu). The distance 
from the nearest locality in Saguache Co. is about 150 miles. 


a 


Voucher specimen: Sandoval Co.: Valles Caldera National Preserve, Jemez Mountains, Cerros del Trasquilar, main 
portion, N35,9648 W106.4918, 8537-8740 ft elev, roadside and coniferous forest, 26 Jun 2001, Hartman 71472 (RM). 


RANUNCULACEAE 

Ranunculus alismifolius Geyer ex Benth. var. montanus S. Watson—According to 
Whittemore (1997), this taxon is restricted to southwestern Colorado, Idaho, Nevada, Utah, 
and Wyoming. It is one of six varieties recognized by this author, all of which occur from 
the Rocky Mountains to the west coast of North America. These specimens document its 
occurrence in New Mexico. 


Voucher specimens: Rio — Co.: San Pedro Mountains, N 36.0667 W106.8433, 10200 ft elev. seep under aspen 
and spruce, 4 Jul 2001 Dorn 8829 (RM); Santa Fe National Forest and Vicinity, Jemez Mountains, Forest Road 70 
and Forest Road 50, Px = as Trail, NW into San Pedro Parks Wilderness to Vacas Trail, Forest Road 51, N36.0421 
W106.7924 to N36.0662 W106.8121, 9300-10135 ft elev, ponderosa pine and some spruce, 20 Jun 2002, Hartman 
75015 (RM); Rito de Abiquiu NE of Cienega del Oso, 2.5 air mi SSE of Polvadera Peak, N36.0290 W106.3900 to 
N36.0360 W106.3820, 9400-10430 ft elev. open riparian bottom and steep shaded slope of corkbark fir/blue 
spruce with an understory of Paxistima, Trauttvetaria, and Vaccinium, : Jul 2003, Reif 6567 (RM); NM hwy 70, 
South San Pedro Parks Wilderness trailhead from parking lot N to San Gregorio Reser voir, 36.0273 N106.8469 to 
W 36.0407 106.8460, 9200-9400 ft elev, open wet meadows in mixed coniferous forest with aspen, 2 Jun 2002, 
Reif 45 (RM 


Ranunculus repens L.—This taxon is naturalized throughout much of the eastern and 
western portions of the United States (Whittemore 1997). The two collections cited be- 
low also appear to represent self-maintaining populations. 


Voucher specimens: Deng al Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, 8 air mi SW of La 
Cueva, Forest Road 376, 2 air mi N of Porter, N35.8442 W106.7810, 7490 ft elev, stream and adjacent meadow, 23 
Jun 2002, Hartman ety (RM). San Miguel Co.: Sangre de Cristo Mountains, Forest Road 60 along Sapello River, 
2 air mi SW of Rociada. N35.8129 W105.4480, 8000 [t elev, canyon bottom peat area at duius a5 trailhead, 
including riparian of river and south-facing bank (below road) of mature aspet Aug 2003, 
Reif 8741(RM) 


ROSACEAE 

Geum triflorum Pursh var. trifloruam—Holmgren (1997) states that var. triflorum ranges 
south in the Rocky Mountains, where it more or less straddles the continental divide 
from Montana to northern New Mexico and includes some populations as far west as 


HARTMAN ET AL., VASCULAR PLANT RECORDS FOR NEW MEXICO 1231 


northeastern Arizona. The following specimens document its occurrence in New Mexico. 
Variety ciliatum, the more western version of the species is listed for New Mexico by Al- 
lred (2005). 


Voucher specimens: Colfax Co.: Philmont Scout Ranch, Beaubien, N36.4237 W105.1033, 9400 - oe common 
on dry, grassy slopes, 27 Jun 1966, Hartman 117 (RM); 6 Jun 1968, Hartman 1819 (RM); Rio A on 
pauona! EOL Canyon Bancosca He air mi W of pace 36.9008 107.2237 to 36.9143 107.2690, Sue: me 7? se 

1-juniper, 28 May 1987, Hartman 22504 with D’Alcamo 
et al. (RM). Sandoval Ce Jemez Mountains, Hack Fore ae River, ca. 0.5 mi above Las Conchas Campground, 
N35.8200 W106.5212, 8500 ft elev, 18 Jun 1986, Jacobs 006 with Jacobs (RM). 


Potentilla fissa Nutt.—This species is common throughout the Rocky Mountains and sev- 
eral adjacent states; likewise it is included for New Mexico (USDA, NRCA 2006). Allred 
(2005) indicates that it is reported for New Mexico but awaits verification. 

Voucher specimen: San Miguel Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Hermit 
Peak, N35.7447 W105.4139, 10000-10212 ft elev, open grassy and rocky areas and rim rock, 18 Aug 2002, Hart- 
man 76734 (RM) 

Prunus persica (L.) Batsch—The introduced peach is known from much of the eastern 
United States and several of the states to the west, but it is not reported from Montana 
south to New Mexico (USDA, NRCS 2006). In both localities here listed, it was found grow- 
ing in sites devoid of human disturbance and thus appears to be adventive. 


Voucher specimens: Sandoval Co.: Valles Caldera National Preserve, Jemez Falls Road to turn south, N35.8277 
W106.6170, 8000-8164 ft elev, ponderosa pine forest with rock outcrops and swales, 30 Jul 2001, Hartman 73177 
(RM); Santa Fe National Forest and Vicinity, Jemez Mountains, Battleship Rock Trail 137 at McCauley Warm 
Springs, N35.8222 W106.6281, 7300-7400 ft elev. warm springs and riparian drainage in os pine forest, 1 
Sep 2003, Reif 9380 (RM). 


SAXIFRAGACEAE 

Heuchera hallii A. Gray—This taxon was considered by Rosendahl et al. 1936) as distinc- 
tive. It has been considered to be endemic to Colorado; in the vicinity of Pikes Peak 
(Chaffee, Custer, Douglas, El Paso, Jefferson, and Park cos.). (http://cumuseum. 
colorado.edu). These reports represent a range extension of about 90 miles to the south. 


ucher — Colfax Co.: Philmont ee Ranch, Rayado Canyon, 0.5 mi W of Old Abreu Lodge, N36.3765 
ae 0331, 7500 ft elev, sandy stream bank and rock Loe 15 Jul 1968, Hartman 2385 (RM); base of Crater 
Peak alon ngR yado River, ca. 2 air mi SE of one Camp, N36.3746 W105.0380, 8200-8800 ft elev, slope above river 
10 Jul 1991, a 92a with Hartman (RM); 4 Aug 1991, a bry 92b(RM). 


SCROPHULARIACEAE 

Penstemon glaber Pursh var. alpinus (Torr.) A. Gray—This variety is the second for the 
species to be documented in New Mexico. Its previous known range was southeastern 
Wyoming to south-central Colorado (Great Plains Flora Association 1986, see key to va- 
rieties). Variety brandegeei (TS. Porter & Rydb.) C. Freeman had been the only taxon of 
this species documented for New Mexico. 


Voucher speci :Los Alamos Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, Mitchell Trail 69, 
from Los Alamos, NW 2.1 air mi to Guaje Reservoir, N35.9310 W106.3432, 7500-8900 ft elev, incised gravel stre- 
ambed through a burn, climbing canyon walls of tuft and rhyolite, toa mixed conifer forest with patchy burns, 
28 Jun 2002, Reif 1379 with Hartman and Jacobs (RM); Taos Co.: Carson National Forest and Vicinity, Sangre de 
Cristo Range, Valle Vidal Unit, ridgeline above corral from Forest Road 1950 toward Little Castilla Peak, N36.7843 
W105.2330, 10,000-12,584 ft - spruce fir and bristlecone pine forest with meadow openings, rocky ridgeline, 
30 Jul 2005, Larson 3373 (RM 


1232 BRIT.ORG/SIDA 22(2) 


ACKNOWLEDGMENTS 
The authors thank the following individuals for verification of specimens: Mary 
Barkworth, Utah State University; Ihsan A. Al-Shehbaz, Missouri Botanical Garden. Also 
thanked are Kelly W. Allred, Guy L. Nesom, and Richard W. Spellenberg for their helpful 
comments on the manuscripts. We thank Jill Larson, M.S. student (RM), for allowing us 
to publish her record of Penstemon glaber var. alpinus. 
REFERENCES 
Aten, C.D. 2004. Ecological patterns and environmental change in the Bandelier landscape. In: T.A. 
Kohler, ed. Village Formation on the Pajarito Plateau, New Mexico: Archaeology of Bandelier Na- 
tional Monument. Univ. of New Mexico Press, Albuquerque. Pp. 19-68. 
Allred, K. 2005 (September). A working index of New Mexico vascular plant names. http:// 
web.nmsu.edu/~kallred/herbweb/ 
Batt, PW. 2002. Carex sect. Phaestoglochin. In: Flora of North America Editorial jeu iaa eds. 2002. 
idae (in part): Cyperaceae. 


Flora of North America north of Mexico.Vol.23.Mag 
Oxford Univ. Press, New York, New York. 

Barkwortn, M.E.,J.McNeit, and J. Maze. 1979. A taxonomic study of Stipa nelsonii (Poaceae) with a key 
distinguishing it from related taxa in western North America. Canad. J. Bot. 57:2539-2553. 

Cronauist, A. 1994.In:A.Cronquist, A.H. Holmgren, N.H. Holmgren, J.L.Reveal, and P.K. Holmgren. Inter- 
mountain Flora. Vascular plants of the Intermountain West, U.S.A. Vol. 5, Asterales. New York Bo- 
tanical Garden, Bronx, New York. 

Furtow, J.J. 1997. Betulaceae. In:Flora of North America Editorial Committee, eds. 1997. Flora of North 
America north of Mexico. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. 
Press, New York, New York. 

Great PLAINS Flora Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence, Kansas 

Hartman, R.L. 1973. Notes. New plant records for New Mexico. SouthW. Nat. 18:241-242. 

Hartman, R.L. 2003. Preliminary checklist of the vascular plants of Valles Caldera National Preserve, 
New Mexico. Distributed by author. 

Hartman, R.L. and B.E. Newson. 2005. General floristic survey of the Valles Caldera National Preserve, 
New Mexico: report on field seasons 2001, 2002, 2003, 2004.Report to the U.S. Geological Survey, 
Biological Resources Division. 

Houmaren, N.H. 1997.In: A. Cronquist, NH. Holmgren,and PK. Holmgren. Intermountain flora. Vascular 
plants of the Intermountain West, U.S.A. Vol. 3, Part A, Subclass Rosidae (except Fabales). New York 
Botanical Garden, Bronx, New York. 

Houmoren, N.H. 2005. In:N.H. Holmgren, PK. Holmgren, and A.Cronquist. Intermountain Flora. Vascular 
plants of the Intermountain West, U.S.A. Vol. 2, Part B, Subclass Dilleniidae. New York Botanical 
Garden, Bronx, New York. 


Jacoss, B.F. 1989. Flora of Bandelier National Monument. Final report: National Park Service. 

Kartesz, J.T. and C.A. Meacoam. 1999. Synthesis of the North American Flora, CD format, Version 1.0. 
North Carolina Botanical Garden, Chapel Hill, North Carolina. 

Naczi, R.F.C. 2002. Carex sect. Deweyanae. In: Flora of North America Editorial Committee, eds. 2002. 
Flora of North America north of Mexico.Vol.23.Magnoliophyta:C linidae (in part): Cyperaceae. 
Oxford Univ. Press, New York, New York. 

Nesom, G.L. 2006. Erigeron. In: Flora of North America Editorial Committee, eds. 2006. Vol. 20. 
Magnoliophyta: Asteridae, part 7: Asteraceae, part 2. Oxford Univ. Press, New York, New York. 

O'Kane, S.L., Jr, D-H. Witken, and R.L. Hartman. 1988. Noteworthy collections: Colorado. Madrono 35: 
72-74. 


HARTMAN ET Al 1233 


Reir, B.P. 2006. A vascular plant inventory of Santa Fe National Forest (including the Valles Caldera 
National Preserve) and vicinity, North-Central New Mexico. M.S.Thesis, Univ.of Wyoming, Laramie, 
Wyoming. 

Reir, B.and R.L. Hartuan. 2005.A floristics inventory of the vascular plants of Santa Fe National Forest 
and the surrounding hydrographic region, New Mexico. Final report: Santa Fe National Forest. 
Rottins, R.C. 1993. The Cruciferae of Continental North America: Systematics of the mustard family 

from the Arctic to Panama. Stanford Univ. Press, Stanford, California. 

RoseNDAHL, C.O., F.K. Butters, and O. Lakria. 1936. A monograph on the genus Heuchera. Minnesota 
Studies in Plant Science, Vol. Il. Univ.of Minnesota Press, Minneapolis 

THeosatp, W.L., C.C. Tsenc, and M.E. MatHias. 1963. A revision of Aletes and Neoparrya (Umbelliferae). 
Brittonia 16:296-315. 

Ureatscu, L.E. and PB. Cox. 2006. In: Flora of North America Editorial Committee, eds. 2006. Vol. 21. 
Magnoliophyta: Asteridae, part 8: Asteraceae, part 3. Oxford Univ. Press, New York, New York. 

USDA, NRCS. 2006.The PLANTS Database, 6 March 2006 (http://plants.usda.gov). National Plant Data 
Center, Baton Rouge, Louisiana 70874-4490. 

Weser, W.A. 1958. Rediscovery of the genus Neoparrya Mathias (Umbelliferae). Rhodora 60:265-271. 

Wetsu, S.L., N.D. Atwoopo, S. GoopricH, and L.C. Hicains (eds.). 2003. A Utah flora. Third edition, revised. 
Brigham Young Univ., Provo, Utah. 

Wuittemore, A.T. 1997. Ranunculus.\|n: Flora of North America Editorial Committee, eds. Flora of North 
America north of Mexico. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. 
Press, New York, New York. 


1234 BRIT.ORG/SIDA 22(2 


Books RECEIVED 


Joun F Leste and Brett A. SUMMFRELL, Photographs by SuZANnr BuLLock. 2000. The Fusarium 
Laboratory Manual. (ISBN 0-8138-1919-9, pbk.). Blackwell Publishing. 2121 State Ave., 
Ames, IA 50014-8300, U.S.A. (Orders: 515-292-0140, 515-292-3348 fax 1-800-862- 
6657, wwwblackwellprofessional.com, orders@ames.blackwellpublishing.com,). 
$124.99, 388 pp., numerous b/w illustrations, 81/2" x 11" 

From the Publisher—‘For the first time in over 20 years, a comprehensive collection of photographs and descrip- 

tions of species in the fungal genus Fusarium is available. This laboratory manual provides an overview of the 

biology of Fusarium and the techniques involved in the isolation, identification and characterization of indi- 
rich they occur. lt is the first time that genetic, morphological and mo- 


vidual species and the populations in w 
pigs oy oaches have been incorporated into a volume devoted to Fusarium identification. The authors in- 
f 


cies, both new and old, and provide protocols for genetic, morphological and molecular 
ide ee techniques. 

“The Fusarium Laboratory Manual also inc 
thinking that has begun to inform the understanding of agriculturally important fungal pathogens. In addition 
to practical “how-to” protocols it also provides guidance in formulating questions and obtaining answers about 


udes some of the evolutionary biology and population genetics 


this very important group of fungi. The need for as many different techniques as possinlet to be saa in the ide n- 
8 ) q 


tification and characterization process has never been greater. The 
than those in the genus Fusarium. This volume presents an eee to the seneris usarium, the toxins retest 


fungi produce and tl the diseases tl they can cause.” 
J 


Kris M. Havstab, LaurA EF HueNneke, and WILLIAM H. SCHLESINGER. (eds.). 2006. Structure and 
Function of a Chihuahuan Desert Ecosystem. The Jornada Basin Long-Term Eco- 
logical Research Site. (SBN 978-0-19-511776-9, hbk.). Oxford University Press, 2001 
Evans Road, Cary, NC 27513, U.S.A. (Orders: 800-451-7556; http://www.oup.com). 
$74.50, 465 pp., b/w halftones, maps and line illustrations, 6 1/8" x 9 1/4". 


From the Publisher—“The Jornada Basin LTER is located in the Chihuahan Desert, the largest in North America. 


This region of south central New Mexico hasa history of nearly 100 years as the basis for scientific research. This 
work gives a thorough, encompassing review of the tremendous array of observations resulting from experi- 
ments conducted in this ecosystem. Beginning with thorough descriptions of the most salient features of the 
region, the book then reviews a wide range of archived and active data sets on a diversity of biotic and abiotic 


features. It next presents a syntheses of important topics including livestock grazing and remediation efforts. A 


concluding chapter provides a synthesis of the principles that have emerged from this body of work, and how 
these relate to the broader fields of ecology and natural resource management. It concludes with recommenda- 
tions for future research directions. The insightful views expressed in this volume should guide management of 
arid landscapes globally. This is the sixth volume in the Long Term Ecological Network Series. 


GALIUM TRICORNUTUM (RUBIACEAE) AND PARENTUCELLIA 
VISCOSA (SCROPHULARIACEAE) NEW TO OKLAHOMA 


Amy K.Buthod Bruce W. Hoagland 
Oklahoma Biological Survey Oklahoma Biological Survey and ie t of Geography 
University of Oklahoma University of Okla 
Norman, Oklahoma 73019, U.S.A. Norman, Oklahoma at és aon 
amybuthod@ou.edu 
ABSTRACT 


This paper reports the occurrence of two species previously unknown to the flora of Oklahoma. Galium 
tricornutum, a Eurasian introduction, was discovered in a cement drainage ditch in Hughes County in central 
Oklahoma. Parentucellia viscosa, an introduction from Europe, was discovered in disturbed grassland in 
McCurtain County in extreme southeastern Oklahoma. 


RESUMEN 
Este articulo cita la ocurrencia de dos — Le tnens de Seon Oclads de a me de ee oa 
tricornutum, una introd iatica, fue descubierta en una zanja d ado de 
Hughes en el centro de Oklahoma. nner ie viscosa (L.) Caruel, una introduccién de Europa, fue descubierta 
en un prado al | >| condado de McCurtain en el extremo sudeste de Oklahom 


| 


Galium tricornutum Dandy, in the Rubiaceae, is an int d annual herb native to Eurasia 
that is typically found in waste places and as a weed of cereal crops. It has been reported 
previously in Arksansas, California, Georgia, New York, Oregon, Pennsylvania, Rhode Is- 
land, South Carolina, Washington, Wisconsin, and Oregon (USDA-NRCS 2006). One plant 
was encountered growing in a crack in a cement drainage ditch in the Holdenville city 
park in central Oklahoma. The combination of recurved pedicels and a tuberculate fruit 
distinguish G. tricornutum from other ey Galium species in Oklahoma. 


Voucher specimen for Galium tricornutum: OKLAH Hughes Co.: Holdenville, at Stroup City Park, T7N R9E 
secl8, ll May 2006, Buthod, Hoagland, and pein a 7138(OKL »: 


Parentucellia viscosa (L.) Caruel, in the Scrc pl 1ulariaceae. is an introduced annual herb na- 


tive to Europe typically found on moist, grassy places and frequently in old meadows. It 
has been reported in Arkansas, California, Hawaii, Louisiana, Oregon, Texas, and Washing- 
ton (USDA-NRCS 2006). Only one plant was located in a disturbed tallgrass prairie site on 
limestone in southeastern Oklahoma. Associated species included Centaurea americana 
Nutt., Echinacea atrorubens, Sorghum halepense (L.) Pers., and Rudbeckia hirta L. 


Voucher specimen for Parentucellia viscosa: OKLAHOMA. McCurtain Co.: 1.9 mi NWof Idabel, T7S R23E SE/4 sec 
25, 25 May 2006, Buthod, Hoagland, and Arbour AB-7137(OKL). 


ACKNOWLEDGMENTS 
We appreciate the field assistance of David Arbour and Priscilla Crawford and the com- 
ments of Susan Barber and Ron Tyrl. 
REFERENCE 


USDA-NRCS 2006. The PLANTS Database [online]. Available: http://plants.usda.gov. National Plant 
Data Center, Baton Rouge, LA 70874-4490 USA. (Accessed on 13 June 2006). 


SIDA 22(2): 1235. 2006 


1236 BRIT.ORG/SIDA 22(2) 


ANNOUNCEMENTS 


2006 Delzie Demaree Travel Award Recipients 


The 18th Annual Delzie Demaree Travel Award was presented at the 52nd Annual Sys- 
tematics Symposium (13-14 Oct 2006) at the Missouri Botanical Garden. Three students 
were presented the Travel Award: Alejandra Vasco, New York Botanical Garden and City 
University; Diana Hurlbut, Central Michigan University; and Michael Sundue, New York 
Botanical Garden and City University. 

The 2006 Travel Awards were underwritten by 1) Delzie Demaree Travel Award En- 
dowment, 2) Members of the Delzie Demaree Travel Award Committee. and 3) John Clay- 
ton Chapter of the Virginia Native Plant Society. 

Anyone interested in making a contribution to Delzie Demaree Endowment Fund, 
which supports the travel award, may make contributions by VISA or MasterCard or by 
a check, payable to Botanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan 
Street, Fort Worth, TX 76102-4060, U.S.A. 1-817-332-7432; Email: barney@brit.org. Thank 
you. 


The 2007 Applications for the Delzie Demaree Travel Award 


Applications for the 2007 Delzie Demaree Travel Award should include a letter from the 
applicant telling how symposium attendance will benefit his/her graduate work and 
letter of recommendation sent by the major professor. Please send letters of application 
to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology Department, The College of 
William and Mary, Williamsburg, VA 23185-8795, U.S.A. 1-757-221-2799; Email: 
ddmware@wm.edu. The period for receiving applications will end three weeks prior to 
the date of the symposium if a sufficient number of applications are in hand at that time. 
Anyone wishing to apply after that date should inquire whether applications are still 
being accepted before applying. The Systematics Symposium dates for 2007 are 12-13 
October 2007. 

The Delzie Demaree Travel Award was established in 1988 and honors Delzie Demaree 
who attended 35 out of a possible 36 symposia before he died in 1987. Delzie Demaree 
was a frontier botanist, explorer, discoverer, and teacher. His teaching career asa botanist 
began in Arkansas at Hendrix College in 1922. He also taught botany at the University of 
Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas AS&tM, and Arkansas 
State University at Jonesboro where he retired as professor emeritus in 1953. One of the 
things he enjoyed most as a botanist was assisting students with their field botany re- 
search. 


SIDA 22(2): 1236. 2006 


CAPRARIA MEXICANA (SCROPHULARIACEAE) 
IN CAMERON COUNTY, TEXAS: 
REDISCOVERED IN THE UNITED STATES 


Alfred Richardson Ken King 
Department of Biological Sciences 1008 W. 5" Street 
The University of Texas at Brownsville Weslaco, Texas 78596, U.S.A. 
80 Fort Brown 


Brownsville, Texas 78520, U.S.A. 


ABSTRACT 


Two new sites are reported for Capraria mexicana (Scrophulariaceae) in the United States. 


RESUMEN 


Se reportan dos nuevos sitios para Capraria mexicana (Scrophulariaceae) en los Estados Unidos. 


Capraria mexicana (Mexican Capraria) is an attractive shrub reaching 2 m in height (Fig. 
1). The leaves are alternate, lanceolate, with serrate margins. The flowers emerge from the 
leaf axils. Corollas are regular, white, basally sympetalous, with lobes about 9 mm long. 
Everitt et al. (2002) provide description and a picture. Most reports of C. mexicana are 
from Tamaulipas, Mexico, with records also from Jalisco, Guerrero, and Tabasco. The 
southernmost collections are from Belize (Ideker 1996). 

apraria mexicana was first discovered in the United States by Joe Ideker in 1993. 
He reported at least 66 mature plants plus seedlings in three small stands located at Fal- 
con State Park near the International Falcon Reservoir in Starr County, Texas. (Ideker 
1996). The plants were in a precarious location where they could be flooded with a rise in 
the reservoir, and they were located within a proposed drilling pad site. After communi- 
cation with Joe Ideker in 1993, this species was listed as Endangered in Texas by the Texas 
Organization for Endangered Species (TOES). It is not considered Endangered in Mexico. 
Our attempt to relocate the population on 25 March 2006 was unsuccessful. Benito Tre- 
vino, well known authority on plants of Starr County, reported having seen no Capraria 
plants when he had visited the site. 

In September of 2005, while at the Laguna Atascosa National Wildlife Refuge, Cam- 
eron County, Texas, we were showna single plant of Capraria mexicana by Ellie Thomp- 
son, volunteer manager of the butterfly garden there. She had watched it grow from a 
volunteer seedling to maturity. As no nearby members of the species were known, it was 
assumed to be an inadvertent introduction from Starr County nursery stock. 

In March of 2006, we observed a large population of Capraria mexicana in 
Brownsville, Cameron County, Texas, along Highway 511, 0.4 miles north of Old Port Isa- 
bel Road. The population continued for at least 0.2 miles. The population is estimated to 
comprise several hundred individuals ranging from 0.3 m to 1.2 m tall The plants were 
seen growing on both sides of the highway, on saline clay, with Opuntia engelmannit var. 
lindheimeri, Borrichia frutescens, Prosopis glandulosa var. glandulosa, P reptans, Maytenus 
phyllanthoides, Fleischmannia incarnata, Dichanthium sp, and Bothriochloa sp. 

With the presumed extirpation of the population at Falcon State Park, the Brownsville 


SIDA 22(2): 1237— 1238. 2006 


1238 BRIT.ORG/SIDA 22(2) 


site is the only confirmed population of Capraria mexicana in the United States. Further 
surveys might reveal a population at Laguna Atascosa National Wildlife Refuge. The 
plants can be easily overlooked if not in flower, since the leaves greatly resemble those of 
Baccharis. 
Voucher specimen: TEXAS. Cameron Co.: Brownsville, Hwy 511, 0.4 mi N of Old Port Isabel Road, 16 Mar 2006, 
Richardson and King 3284 (BRIT, TEX). 
ACKNOWLEDGMENTS 
We thank Mike Heep for identification of the grasses, and an anonymous reviewer who 
provided useful and constructive comments. 
REFERENCES 
Everitt, J., D.L. Drawe, and R.I. Lonarp. 2002. Trees, shrubs & cacti of South Texas. Second Edition. Texas 
Tech University Press, Lubbock. 
IbeKer, J. 1996, Capraria mexicana (Scrophulariaceae), an endangered addition to the United States 
Flora.. Sida 17:523-526 
TEXAS ORGANIZATION FOR ENDANGERED Species (TOES). 1993.Endangered, threatened and watch lists of Texas 
plants. TOES Publ. 9, a rev (and 1993 addenda). 


RANGE EXPANSION OF HEXALECTRIS GRANDIFLORA 


(ORCHIDACEAE) IN TEXAS 
el Brown-Marsden Anne B. Collins 
iolo partment a of Land, Air and Water Resources 
Dee of Dallas ls and Biogeachemistry 
1845 F. Northgate Drive Fes of California at Davis 
Irving, Texas 75062, U.S.A. 09 Veihmeyer Hall 


Davis, California 95616, U.S.A. 


ABSTRACT 


Plants of Hexale liflora A. Richard & Galeotti (Orchidaceae) collected in Dallas County, Texas in 2005 


and 2006 ey fie oe of this species northeast by more than 700 km and indicates greater diversity for the 
cas enus He ke in ee Coun than previously observed. These observations suggest either the absence of 


t data on species’ range in intervening counties, or the loss of populations between the north central 
Texas and western Tees locations, and argue for continued study of Hexalectris throughout the state. 


ABSTRACT 
Las plantas de mealecin isgra ayn Bs Richard & Sai (Orchidaceae) colectadas en el condado de Dallas, 
ana en 2) v2 | noreste mas de 700 km e indican mayor diversidad 
lectri | lado de Dallas de la q Estas observaciones 
l iaded fici del Ide est ie en los eondndcs afectados, o la pérdida de 


pencones entre el norte-centro de Texas y las local idades del oeste de Texas, y se aportan argumentos para 
continuar el estudio de Hexalectris en todo el estado. 


The myco-heterotrophic orchid genus Hexalectris Rafinesque is comprised of seven spe- 
cies—five that are found in the United States and two that are found only in Mexico (Gold- 
man et al. 2002). Four of the species found in Texas are H. grandiflora A. Richard & 
Galeotti, H. warnockii Ames & Correll, H. nitida L.O. Williams, and H. revoluta var. revoluta 
Correll (H. revoluta var. colemanii Catling) has recently been described from Arizona 
(Catling 2004). The fifth species found in Texas, H. spicata (Walter) Barnhart, includes 
two varieties: H. spicata var. spicata and H. spicata var. arizonica (S. Watson) Catling & 
Engel . Hexalectris spicata var. spicata. ranges widely from Arizona through the south- 
eastern US and north to Maryland, and H. spicata var. arizonica, revalidated in 1993, is 
found only in parts of Texas and Arizona (Catling & Engel 1993). 

Dallas County has recorded populations of H. spicata var. spicata, H. spicata var. 
arizonica, H. warnockii,and H. nitida (Diggs et al. 1999; Hatch et al. 1990). Dallas County 
populations of Hexalectris are found in areas of the White Rock Escarpment, a geologi- 
cal region of lower Cretaceous limestone that extends northeast from Dallas to the Okla- 
homa border, and southwest past Waco and Austin into the Edwards Plateau (Dallas 
Department of Urban Planning 1977). The escarpment geological zone, known for its dis- 
tinctive flora, is most evident in the southwestern part of the county, and it is in this area 
where most of the Dallas County plants of Hexalectris have been found (Collins et al. 
2005). Studies in Dallas County examined habitat structure and soil association for 
Hexalectris. They indicate an association with oak litter (generally Quercus buckleyit or 
Q. shumardii) mixed with Juniperus spp. and distributed primarily on Dallas soil series 
Eddy-Brackett complex (8-20 % slope) (Collins et al. 2005). Ongoing work attempts to 


SIDA 22(2): 1239- 1244. 2006 


1240 BRIT.ORG/SIDA 22 


= 


2) 


confirm this soil-orchid relationship through targeted searches in preserved habitat ar- 
eas with Eddy-Brackett soils that are well-drained and have sufficient slope. 

Intensive studies of Hexalectris distribution in 2005 and 2006 led to the discovery 
of H. grandiflora, a species not previously found in Dallas County. On 21 June 2005, dur- 
ing Hexalectris studies on the focal soil series,a cluster of H. grandiflora was identified at 

Camp Wisdom, a 344-acre tract of escarpment habitat in southwest Dallas County owned 
and operated by the Circle 10 Council of Boy Scouts of America as an educational and 

See site. One cluster of six aboveground stems was found during a comprehensive 
search of the location (Fig. Ll). The stems were erect, leafless, and slender, pinkish red at 
base, and becoming lighter toward the inflorescence. The dorsal sepals were oblong to 
elliptic; white to pale pink at base, and light pink distally. The lateral sepals were falcate, 
white to pale pink at base and light pink distally. The petals were recurved, white to pale 
pink at base and light pink distally. The column was pink with a white tip and the lip 
(labellum) was lobed and lamellate. The lateral lobes were deep pink, and the middle 
lobe was pink at the margin and white at the center. Stems were found in an area domi- 
nated by Quercus spp. and Juniperus spp., although the plants were found directly below 
arusty blackhaw viburnum (Viburnum rufidulum). Fifteen blooming stems were noted 
on 9 June 2006 at the same location, and were observed to be more robust than when first 
sighted in 2005. 

A second population of Hexalectris grandiflora was found at Cedar Ridge Preserve 
in 2006 approximately 5.5 km southwest of the 2005 sighting. A cluster of 9 stems was 
observed on 13 May 2006. Definitive identification of the species was possible on 2 June 
2006 when the plants were blooming (Fig. 2). These Hexalectris stems were immediately 
adjacent to one of the preserve’s trails in an area dominated by Quercus spp. and Jun iperus 
spp, and with a somewhat open canopy due toa trail cut. Only half of the individuals 
survived to flowering, due to losses from trail use and inadvertent trampling of the small 
stems. 

Specimens (Fig. 3) from the Camp Wisdom and Cedar Ridge locations were depos- 
ited at the herbarium at the Botanical Research Institute of Texas in Fort Worth (BRIT). 
In addition, tissue from the plant stem and roots was sampled at both locations for ge- 
netic analysis as part of a study of phylogeny, evolution and mycorrhizal specificity in 
Hexalectris orchids (A. Kennedy, pers. comm.) 


Voucher specimens: Texas. Dallas Co.: Camp Wisdom (6400 Redbird Ln.), Escarpment habitat, Eddy-Brackett 
soil series, N 32 40.552’ @ 96 56.640, elev. 730 ft, 21 Jun 2005, M. Brown-Marsden and A. Collins s.n. (BRIT): Cedar 


Ridge Preserve, White Rock Escarpment, Dallas Park and Recrreation/Dallas County Open a property, N 32° 
37.056 W 096’, elev. 711 ft, mixed oak juniper woods, ground cover primarily juniper and oak leaves, some bare 


ial 


rock, adjacent to trail, one of nine stems seen emerging on 13 Mz vy 2006, first blooms 2 Jun 2006, M. Brown-Marsden 


sn. (BRIT) 


In Texas, Hexalectrisgrandiflora has previously been documented only within the Davis 
and Chisos Mountains of extreme western Texas at elevations of 700-2500 m (Goldman 
et al. 2002). Confirmation of H. grandiflora in Dallas County marks both a significant 
disjunct in its distribution, and deviation from previously described habitat types. The 
plants found at the Camp Wisdom and Cedar Ridge sites were at elevations of 217 and 
243 m, respectively. They were found in woodlands dominated by oak and juniper, but 
lacking the pinyon pine of the extreme west Texas part of its normal range. In spite of 
these deviations from the range and habitat of conspecific populations, they were still 
found on the same soil types as other members of the genus throughout Dallas County 


BROWN 1 AND COLLINS TEXAS 1241 


lam 
ie 


Fic. 1, Photograph of Hexalectri liffora infl t Camp Wisdom, Dallas County, Texas on 21 June 2005. Photo by Randy 
Schoen. 


(Collins et al. 2005). Additionally, all observations of H. grandiflora have occurred within 
years of moderate to severe drought, and have been the driest years on record since 1963- 
1964 (Drought Preparedness Council report, March 2006). 

The discovery of this significant disjunct in the range of Hexalectrisgrandiflora gives 
rise to larger questions with broader conservation implications. If this species is present 
in Dallas County but unknown in the area between Dallas and its previously known 
range, the gap may result from a lack of studies on Hexalectris ecology and distribution. 
In this event, it would be possible for this species to be found in one or more of the inter- 
vening counties through targeted studies such as those conducted in Dallas County. How- 
ever, it is also possible that the historic range of H. grandiflora included area between 


1242 BRIT.ORG/SIDA 22(2) 


Fig, 2, Photograph of Hexalectris grandiflora inflorescence at Cedar Ridge Preserve, Dallas County, Texas on 2 June 2006. Photo by 
Puttaswamy Ravishankar. 


Dallas and West Texas, but this has since declined due to increased urbanization, and 
habitat loss and degradation. Determination of the ultimate causes and consequences of 
the recorded range disjunct may depend on continuing to answer fundamental ecologi- 
cal questions about this genus and factors influencing persistence of Hexalectris th rough- 
out its range. 


BROWN LLIN 1243 


260 Kilometers 


Fic. 3. Map of historic locations of Hexalectris grandiflora in Texas (circles; after Catling 2004) and the new locations found in Dallas 
County (triangle). 


ACKNOWLEDGMENTS 


We thank Circle 10 Council Boy Scouts of America admini ion and staff for allowing 
us access to the Camp Wisdom site. In particular, we acknowledge the assistance of Randy 
Schoen during our initial censusing effort, and Phil Augsberger, camp ranger at Camp 
Wisdom. Jim Varnum, Stephanie Varnum, Rigel Rilling, and Jenny Krabacher provided 
assistance in fieldwork. Aaron Kennedy (Miami University-Ohio) assisted in collecting 
tissue for laboratory analysis. Photography assistance was provided by P. Ravishankar. 


REFERENCES 


Carunc, BM. 2004. A synopsis of the genus Hexalectris in the United States and a new variety of 
Hexalectris revoluta. Native Orchid Conf. J. 1(2):5-25. 

Carine, PM. and VS. Encet. 1993. Systematics and distribution of Hexalectris spicata var. arizonica 
(Orchidaceae). Lindleyana 8:119-125. 


1244 BRIT.ORG/SIDA 22(2) 


Cottey, J. 2006. Statewide drought situation report. Report of the Drought Preparedness Council, 
March 6, 2006. 

CoLuns, A., J. Varnum, and M. Brown-Marspen. 2005. Soil and ecological features of Hexalectris 
(Orchidaceae) sites. Sida 21:1879-1891. 

DALLAS DEPARTMENT OF UrBAN PLANNING. 1977. The escarpment report. Department of Urban Planning of 
the City of Dallas, Texas. 

Diccs, G.M., Jr., B.L. Liescome, and R.J. O’KENNON. 1999, Shinners & Mahler's llustrated Flora of North 
Central Texas. Botanical Research Institute of Texas, Fort Worth. 

Gotoman, D.H., R.A. Coteman, L.K. Macrath, and PM. Cattinc. 2002. Hexalectris. In Flora of North America, 
Vol. 26. Oxford University Press, New York and Oxford. Pp. 603-607. 

Hatcu, S.L., K.N. GaNoni, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. 
Sta. Misc. Publ. 1655:232-235. 

Licaio, J.and A.O. Liccio. 1999, Wild Orchids of Texas. University of Texas Press, Austin, 


ERODIUM MALACOIDES (GERANIACEAE), NEW TO TEXAS 
David E.Lemke and Jed L.Aplaca 


Department of Biology 
Texas State University 
San Marcos, Texas 78666-4605, U.S.A. 


ABSTRACT 
Erodium malacoides (Geraniaceae) is reported as an addition to the flora of Texas. A description of the species 
and a key for identifying the Te pecies of Erodium are provided. 

RESUMEN 


Se cita Erodium | id i ) nueva adicion a la flora de Texas. Se aporta descripcion 


de la especie y una clave de las especies de Erodium de Texas. 


Erodium L'Hér. comprises ca. 60 species of annual or biennial, rosette-forming herbs wide- 
spread in temperate and subtropical regions of the Old World and New World (Mabberley 
1997). Three species have previously been recorded from Texas: the native FE. texanum A. 
Gray, which is broadly distributed across central and southern portions of the state west 
to Utah and California, and the non-native species E. cicutarium (L.) LHér. and E. botrys 
(Cav. Bert. Jones et al. 1997). In Texas, E.cicutarium occurs ina variety of disturbed habi- 
tats in central, northern, and western regions of the state, while E. botrys has been col- 
lected from similar habitats in Bexar, Taylor, and Throckmorton counties (Correll & 
Johnston 1970; Turner et al. 2003). 

In the spring of 2005, the senior author discovered a small population of Erodium 
malacoides(L.) Willd, a species previously unknown from the state, established on a dis- 
turbed site in Hays County, Texas. By 2006 the population had increased in size to hun- 
dreds of individual plants and occupied a substantially larger area, indicating a poten- 
tial for naturalization. In the system proposed by Pysek et al. (2004), FE. malacoides would 
be categorized as a “casual alien” in Texas. 

Voucher specimens: U.S.A. TEXAS. Hays Co.: E side of northbound frontage road of Interstate Hwy. 35, City of 
San Marcos, N29° 53’ 38", W97° 54' 31", 02 May 2006, Aplaca 287 (SWT): W side of northbound frontage road of 
Interstate Hwy. 35, City of San Marcos, N29° 53' 39", W97° 54’ 31", 02 May 2006, Aplaca 288 (SWT), E side of 
northbound frontage road of Interstate Hwy. 35, City of San Marcos, N29° 53’ 45", W97° 54' 25", 02 May 2006, 
Aplaca 289 (SWT); 

We can only speculate as to how E. malacoides may have been introduced at this loca- 
tion. Several years ago, activity associated with the construction of a new entrance ramp 
onto the interstate highway disturbed portions of the area now occupied by the species. 
The seed mix used for revegetation of the highway right-of-way, soil brought in as fill 
material, or vehicular traffic from Mexico represent the most plausible seed sources. 

Erodium malacoides is native to southern Europe (Brandes 1995) but has become 
naturalized across the Mediterranean region (El Hadidi et al. 1984). It has been intro- 
duced as widely as Australia and New Zealand (Webb etal. 1988; Westbrooke & Florentine 
2005), South America (Figueroa et al. 2004), and North America, where it has been re- 
ported from a number of Mexican states (Villasenor & Espinosa-Garcia 2004) and from 


SIDA 22(2): 1245- 1249. 2006 


BRIT.ORG/SIDA 22 


Fic. 1. Upper photo: Individual plant of Erodium malacoides from Hays Co., Texas, showing the cha istic simple, palmately lobed 
leaf and relatively small (ca. 10 mm diam.) flower. Lower phowo: Close-up of flower showing the glandular pubescence of the calyx 
and the broad, entire-margined staminodes. 


LEMKE AND APLACA 1247 


California (Taylor 1993), New York (Mitchell & Tucker 1997) and Massachusetts (Magee 
& Ahles 1999) in the U.S.A. 

Plants of E. malacoides (Fig. 1) are annual herbs with decumbent to ascending, pu- 
berulent stems that are glandular at the nodes. The leaves are simple, 4-15 cm long in- 
cluding the petiole, which is generally longer than the blade. The leaf blades are ovate, 
crenate to shallowly palmately lobed, and puberulent. The small flowers are borne in 
umbellate clusters and have glandular-pubescent sepals 4-6 mm long, lavender to purple 
petals about as long as the sepals, five fertile stamens, and five broad, entire-margined 
staminodes. The fruit is a five-parted schizocarp that separates from the style column 
into single-seeded segments, each consisting of an indehiscent fusiform body witha more 
or less rounded, glandular apical pit subtended by a single furrow, and a persistent style 
segment that becomes tightly coiled when dry. The following key is provided to distin- 
guish among the four species of Erodium presently known from Texas. 


KEY TO THE SPECIES OF ERODIUM IN TEXAS 


1. Leaf blades simple, often shallowly or deeply palmately lobed 
2. Stems appressed-canescent; sepals 5-10 mm long, apically strigose; petals unequal, 7-12 
m long; staminodes long-awned, conspicuously ciliate; fruit body 5-8 mm long, the pit 
ae elliptic and eglandular, furrows absent; style column 3-7 cm long E.texanum 
2. Stems puberulent, the nodes glandular; sepals 4-6 mm long, glandular; petals equal, 4-6 
mm long; staminodes awnless, entire-margined; fruit body 3-5 mm long, the pit + round, 
glandular, and usually subtended by one furrow; style column 2-3 cm long E. malacoides 
]. Leaf blades deeply pinnately lobed to pinnately compound 
3. Stem pubescence glandular; sepals 3—5 mm lonag;fruit body 4-7 mm long, the pit subtended 


by a single furrow or furrow absent; style column 2-5 cm long E. cicutarium 
3. Stem pubescence eglandular; sepals 10-13 mm long; fruit body 8-11 mm long, the pit usu- 
ally subtended by two furrows; style column 5-12 cm long E. botrys 
REFERENCES 


Branpes, D. 1995. The flora of old town centres in Europe. In: Sukopp, H.,M. Numata and A. Huber, 
Urban ecology as the basis of urban planning. SPB Academic Publishing by, Amsterdam, The 
Netherlands. Pp. 49-58. 

Corrett, D.S.and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Founda- 
tion, Renner. 

Et Haoiol, M.N., A.A. Faveo, and S.M. Et Nacaar. 1984. Systematic revision of Erodium (Geraniaceae) in 
Egypt. Pl. Syst. Evol. 144:307-314. 

FicueroA, J.A., S.A. Castro, PA. Marauet, and FM. Jaksic. 2004. Exotic plant invasions to the Mediterranean 
region of Chile: causes, history and impacts. Revista Chilena Hist. Nat. 77:465-483. 

Jones, S.D., J.K. Wiprr, and P.M. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist 
including synonymy, bibliography and index. Univ. of Texas Press, Austin. 

Massertey, DJ. 1997. The plant-book. A portable dictionary of the higher plants, 2nd ed. Cambridge 
Univ. Press, Cambridge. 

Macce, D.W.AND H.E. Antes. 1999. Flora of the Northeast.A manual of the vascular flora of New England 
and adjacent New York. Univ. of Massachusetts Press, Amherst. 

MitcHett, R.S.and G.C. Tucker. 1997. Revised checklist of New York State plants. Contributions to a flora 
of New York State, checklist IV. New York State Mus. Bull. 790:1-400. 

Pysek, P, D.M. RicHarpson, M. RemAnek, G.L. Weester, M. WILLIAMSON, and J. KirscHNer. 2004. Alien plants in 
checklists and floras: towards better communication between taxonomists and ecologists. Taxon 
53:131-143 


1248 BRIT.ORG/SIDA 22(2) 


Taytor, M.S. 1993. Erodium. In: Hickman, J.C.,ed. The Jepson manual: higher plants of California. Univ. 
of California Press, Berkeley. Pp. 672-673. 

Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003.Atlas of the vascular plants of Texas. Sida, Bot. Misc 
24:1-888. 

Vittasenor, J.L. and FJ. Espinosa-Garcia. 2004. The alien flowering plants of Mexico. Diversity Distrib. 
10:113-123 

Wees, C.J., WAR. Sykes, and PJ. Garnock-Jones. 1988. Flora of New Zealand IV: naturalized dicots, gymno- 
sperms, ferns and fern allies. Manaaki Whenua Press, Lincoln, New Zealand. 

Wesrerookt, M.E. and S.K. Florentine. 2005.Rainfall-driven episodic flood events: are they a major factor 
in moulding New South Wales arid land vegetation patterns? Austral. Geogr. 36:171-181. 


NEW DISTRIBUTION RECORDS FOR EUPATORIEAE (ASTERACEAE) 
INTHE UNEEE DS TALES 


Guy L.Nesom Billie L. Turner 
Botanical Research Institute of Texas Plant Resources Center 
509 Pecan Street University of Texas 
Fort Worth, Texas 76102-4060, U.S.A. Austin, oe 78713, U.S.A. 
ABSTRACT 


Several species of BupAtoneae a occur in neoutnenn Arizona and New Mexico at the northern end of their otherwise 


Mexican distribution: St m New Mexico by a single collection, the exact locality un- 


known and perhaps in medal advicent ve Ageratina (Eupatorium) thyrsiflora (known from Ari- 
zona by a single ae and Koanophyllon (Eupatorium) palmeri (previously confused with K. 
solidaginifolium). Re of K. palmeri or K. solidaginifolium from New Mexico have not been confirmed. 
Chromolaena(Eupator ) bigelovii, primarily a species of northeastern Mexico, is d d for the U.S.A. by 
three Texas collections—its occurrence was recorded in Arizona (cited from only the type collection), but that 
record apparently reflects a confusion of label data for the type, which almost certainly was collected along the 
Texas-Mexico border. Lectotypes are chosen for Eupatorium arborescens, Koanophyllon (Eupatorium) palmeri 


(var. palmeri), and Ageratina (Kyrstenia) thyrsiflora. 


RESUMEN 


Vanigs especies de Eupatorieae viven en el sur de Arizona y N 
a pe ee ‘ ray eee pe | 


] de NI Mévi la recoleccién 


7) 
cuya exacta localidad esd i | México), A ti E tori ) th ifl 


de Arizona con una sola leans y Koanophyllon (Eupatorium) palmeri Gone contindic con K. 
solidaginifolium). Las citas de K. palmeri o K. solidaginifolium de Nuevo México no han sido confirmadas. 
Chromolaena (Eupatorium) bigelovii, primariamente una especie del noreste de México, se documenta para los 
Estados Unidos mediante tres colecciones de Texas—su ocurrencia os citada en Arizona (timicamente don la 


coleccién del tipo), p sa cita refleja nfu e los datos de la etiqueta del tipo, que casi 
con seguridad fue colectado alo largo de la tronitera de Texas y México. Se eligen lectotipos de coauall 
arborescens, I } Eup ium) palm var. palmeri), y Ageratina (Kyrstenia) thyrsiflor 


Preparation of Eupatorieae treatments for Mexico and the Flora of North America (FNA) 
region has brought to light three species previously unreported for the United States. Both 
are primarily distributed in western Mexico and are newly recognized as occurring in 
Arizona and New Mexico. A third primarily Mexican species is rare in Texas; it previ- 
ously has been recorded as a member of the Arizona flora but, as discussed here, it does 
not occur in that state. Morphological descriptions of all four species are provided in the 
FNA treatments. 


1. Stevia salicifolia 

This species apparently has not been included for the U.S.A. in any published floristic 
account, but it was recorded in a dissertation study (Grashoff 1972) from southwestern 
New Mexico by a single collection of E.A. Mearns, naturalist for the United States and 
Mexican Boundary Survey of 1892-1894. Grashoff cited the collection as from Grant 
County, but the San Luis Mountains are a range primarily of northern Sonora and Chi- 
huahua, Mexico, with an extension into Hidalgo County. 


Voucher specimen: U.S.A. New Mexico. [Hidalgo Co.]: canon east side of San Luis Mts., 11 Sep 1893, Mearns 2220 
(US) 


SIDA 22(2): 1249- 1253. 2006 


1250 BRIT.ORG/SIDA 22(2) 


In further refinement of the San Luis Mountains locality, Hubbard (1999) concluded that 
Penstemoncampanulatus, which had been considered to be represented in the U.S.A. flora 
only by a Mearns collection, was collected on the Mexican side of the border rather than 
in New Mexico (canon, east side San Luis Mts., 11 Sep 1893, Mearns 2222 - US). In view of 
this, the U.S.A. record documented here for Stevia salicifolia should be considered doubt- 
ful, even though it seems likely that the species has stepped across the border in one place 
or another. 

Geographic-ecological summary.—Flowering Jul-)Aug-Oct (in northern Mexico). 
Oak and oak-pine woodland, rocky sites, crevices, boulder pockets; ca. 1500-2650 m (data 
from northern Mexico). New Mexico; Mexico (Michoacan, Edo. Mexico, Puebla, and Ver- 
acruz northward to Sonora, Chihuahua, Coahuila, Nuevo Leon, Durango, Zacatecas, San 
Luis Potosi). The range of var. salicifolia also closely approaches Texas and Arizona (see 
map in Turner 1997). 

Stevia salicifoliais a variable species—many infraspecific taxa have been named but 
most were regarded by Grashoff as synonyms of var. salicifolia. Among the U.S.A. species 
of Stevia, S. salicifolia is distinct in its glabrous and shiny stem and leaf surfaces, which 
results from the viscid exudate of sunken glands; stems and leaves of the other U.S.A. 
species usually are sparsely pubescent and not ee 
Stevia salic ifolia Cav le 32. pl. 354.1797. “The tyre... came from plants cultivated at the Royal Botani- 

cal Garden, Madrid; Ortega states that the seed was sent hon Mexico by Sesse.” (Mc Vaugh 1984, p. 895). 


2. Ageratina thyrsiflora 

This species is native primarily to the states of Sonora, Chihuahua, Sinaloa, Durango, 
and Jalisco in western Mexico (Mc Vaugh 1984; Turner 1997). It is known from the United 
States only by a single collection near the international border. 

U.S.A. Arizona. [Santa Cruz Co.|: near Nogales, 15 Sep 1929, Harrison & Kearney 6039 (ARIZ fide Phil Jenkins, LL). 
Geographic-ecological summary.—Flowering Sep-Nov. Rocky sites, oak woodland; 1000- 
2200 m (data from Mexican collections at TEX-LL and published notes of McVaugh 1984). 
Arizona; Mexico (Sonora, Chihuahua, Sinaloa, Durango, Jalisco). 

Ageratina thyrsiflora is recognized by its strict, unbranched or few-branched stems 
with alternate, densely arranged leaves, small heads densely clustered in a single, termi- 
nal aggregate, prominently orange-veined corollas and phyllaries, and close cauline and 
petiolar puberulence of minute, sharply upwardly bent hairs. Morphology is relatively 
uniform over the geographic range, except in two collections from Sonora (Muller 3655, 
LL) and Chihuahua (Gentry et al. 18026, ARIZ, LL), where stipitate glands are mixed with 
the eglandular puberulence on lower stems and petioles. One collection from Jalisco 
(Mc Vaugh 21771, LL) also shows a tendency to produce stipitate glands. 

Ageratina thyrsiflora (E. Greene) R. King & H. Robinson, Phytologia 19:227.1970. Kyrstenia 
hyrsiflora E. Greene, Leaf. Bot. Observ. Crit. 1:9. 1903. Eupatorium thyrsiflorum (E. Greene) B.L. Robin- 


son, Proc. Amer. Acad. Arts 43:36. 1907. Type: MEXICO. CHIHUAHUA: Southwestern Chihuahua, Aug 1885, 

i, ae 275 (LECTOTYPE selected here: US, internet image!). In the protologue, Greene cited the following: 
Chihuahua, Mexico, chiefly southward in the State; aes oy Palmer, P ae und E.A. Goldman, and 
lways distributed for E. occidentale var. Arizonicum. t GH was cited by Mc Vaugh (1984) 


as “isotype” of K. thyrsiflora (near C ce 1 Oct 1885, CG. Pringle 613), but there is no duplicate of 
this at NDG (fide Barbara Hellenthal) or at US. Greene perhaps studied the Palmer specimen at US before 
his time at the Smithsonian as an associate in botany (in 1904-1909); he did not annotate ses ees 


howe, n though he did make handwritten notations on U sheets of K 


sheet 


K. laeta, K ne ee and kK. rufa (fide Barbara Hellenthal, pers. comm), 


A 1251 


Eupatorium thyrsiflorum var. holoclerum B.L. Robinson, Proc. Amer. Acad. Arts 43:36. 1907. 
Type: MEXICO. DURANGO: City of Durango and vicinity, Apr-Nov 1896, E. Palmer 755 (HOLOTYPE: GH; 
IsoTYPEs: MO|, US internet image!). 

3. Koanophyllon palmeri 

Koanophyllon (Eupatorium) palmeri is native primarily to western Mexico (McVaugh 

1984; Turner 1997) and is known from the U.S.A. by collections from south-central Ari- 

zona. A similar taxon (Eupatorium palmeri var. tonsum B.L. Robinson) occurs in near- 

coastal localities from Michoacan to Colima, Jalisco, Nayarit, and Sinaloa. The Arizona 
plants are var. palmeri( hoccurs southward through Sinaloa, Chihuahua, and Sonora 

to Durango and Jalisco). Eupatorium arborescens (as cited below) from Chihuahua is a 

synonym of K. palmeri var. palmeri. 


Vouchers for records of Koanophyllon palmeri in southwestern New Mexico (Hidalgo and 
Grant counties—as mapped by Martin and Hutchins 1980) have not been located in New 
Mexico herbaria J. Mygatt, UNM, pers. comm. and R. Spellenberg, NMC, pers. comm.). 
The basis for the Hidalgo County record may bea citation by Wooton and Standley (1915, 
p.647) of a Charles Wright collection “Guadalupe Pass, Wright 1146,” [4-5 October, 1851). 
According to Wooton and Standley (1915, p. 647), “Guadalupe Pass is on the southern 
boundary of the State, and Wright’s specimens may have come from either Mexico or 
New Mexico.” It seems reasonable to maintain K. palmeri asa member of the New Mexico 
flora, at least tentatively, especially in view of the map points shown by Martin and 
Hutchins (1980). 
U.S.A. Arizona. Cochise Co.: Guadaloupe Mts., Guadaloupe Canyon, 13 Oct 1946, Darrow et al. 3562 (LL). Pima 
o.: W side of Mt. Baboquivari, ees 1944, Gould ce 2670 (LL); Baboquivari Mts., 25 Sep, 1927, Harrison 4754 
(LL); Baboquivari Mts., Sep 193] 932 (LL); Sta. Catalina Mts., Sabino Canyon, 27 Sep 1934, deed 
Peebles 10257 (LL); Babocunee Mts., 30 Sep 1934, Kearney & Peebles 10373 (LL); Baboquivari Canyon, 1] Oct 1 
Peebles et al. 413 (LL); Tucson Mts., Hugh Norris Trail, 7 Nov 1976, Urry 840 (TEX), Waterman Mountains, NE = 
of road to Silver Hill Mine, 5 Dec 2001, Van Devender 2001-1014 (TEX). Santa Cruz Co.: Coronado Natl. Forest, just 
to the E end of Forest Rd 217, near the Dos Amigos Mine, E of of Old Glory Canyon and Warsaw Canyon, 31 Aug 
2001, Goldman 2108 (BRIT); Peck Canyon, Atascosa Ranch, 18 Nov 1981, Van Devender s.n. (TEX): ene Canna, 
lmi NW of Bartlett Mts., 29 Oct 1981, Van Devender & Toolin s.n. (TEX). 


Geographic-ecological summary.—Flowering Sep-Nov(-Dec). Shaded rocks along streams, 

crevices, often in oak woodland; 850-1500 m. (data from Arizona collections). 

ie eae palmeri (A. Gray) R. King & H. Robinson, Phytologia 22:150. 1971. Eupato- 
d 


m palmeri A. Gray, Proc. Amer. Acad. Arts 21: 383.1886. Gray did not cite a aaa collection but note 


“type locality, ‘shady places high up in mountains above Batopilas.” Type: MEXICO. [CHIHUAHUAl: South- 

western Chihuahua, August to N ber 1885, Palmer 144 (LECTOTYPE ee ere: GH}; ISOLECTOTYPE: 

US internet image!). Ape collection, E. Palmer 263 (GH)), is similarly labeled. Both GH sheets were 

eee oa ie as “Eupatorium palmeri n. sp.” McVaugh’s account (1956) of Palmer’s seh indi- 
the vicinity of Batopilas were Aad in August through early October 1 


Eupatorium arborescens M.E. Jones, Contr. West. Bot. 12:43. 1908. Type: MEXICO. CHIHUAHUA: 
Sierra Madre Mts., Guayanopa Canon, 3600 feet alt., in the Tropical Life Zone, 24 Sep 1903, M.E. Jones s.n. 
(LECTOTYPE pace ene ne ea hoe 1799, phOscops: ISOLECTOTYPE: RSA-POM 41800, photocopy!). 
Hi | g label oa and probably are ua: of Ene same Gels ec- 
tion; 41799 b gl I h: 41800 | } Each sheet is labeled 

apparently that of f Philli ip Munz, fide Michael Denslow at RSA-POM) “Part of type, Eupatorium ar horescens 
fos ’ Leaf position, vestiture and involucral features are those of K. palmeri, although the acuminate 
leaf apices are more similar to those of K. solidaginifolium. 


a: 


Plants of Koanophyllon palmeri in Arizona have been previously identified as K. 
solidaginifolium (A. Gray) King & H. Robinson (= Eupatorium solidaginifolium A. Gray, 


1252 BRIT.ORG/SIDA 22(2) 


see citation below). As interpreted here, however, and in Turner (1997) and Blake (1924), 
K. solidaginifolium occurs in north-central Mexico (eastern Chihuahua, Coahuila, north- 
eastern Durango, northern Zacatecas) and into the trans-Pecos area of Texas, east of the 
range of K. palmeri. Their ranges approach each other in central Chihuahua, but the two 
apparently are allopatric. The following contrast separates the two species in the U.S.A. 
and northwestern Mexico. 
Leaves all opposite, blades apically acute, upper surfaces sparsely strigose to weakly hispidulous; 

involucres (3—)3.5-4 mm long; phyllaries herbaceous to the margins, at least the outer sald U 

lent, all usually weakly ciliate, outermost ovate-elliptic to obovate nophyllon palmeri 
Leaves usually subopposite to alternate on distal third of stem, blades apically long-acur serine 

upper surfaces glabrous; involucres 4.5—5.5 mm long; phyllaries with narrow but distinct t bas 

line-translucent margins, glabrous and eciliate, outermost narrowly lanceolate. phyllon 

side aici 


Eupatorium ue ium A. Gray, Smithsonian Contr. Knowl. 3(5) [Plant. Wright. 1]: 
ou paloie ye -“Collected in Expedition from Western Texas to El Paso, New Mexico, May-October, 1849, 
by Charles Gey [as on label], Wright 256 (HOLOTYPE: GH, photocopy! lsoTYPE: GH, photocopy!). Gray’s 
citation in the protologue was “Mountains between the Limpia and the Rio Grande, New Mexico.” Ac- 

ield notes (fide Walter Kittredge, GH), this locality 


cording to LM. Johnston’s commentary on Wright's fi 
matches Wright ’s field number of 256, as recorded on the type labels. L ee creek and Limpia Canyon 
are in heen day Jeff Davis Co., Texas, and it is probable that Wright's collection was made there (recent 


t the species in Jeff Davis Co. and Presidio Co.), proba bly in late August (as inferred 


from Wooton 1906). The specimens were annotated as ‘holotype’ and ‘isotype’ by D.E. Boufford in 1981. 


4. Chromolaena bigelovii 
Chromolaena bigelovii has been included in various summaries of the Arizona flora (e.g, 
Kearney & Peebles 1951; Lehr 1978) on the basis of the protologue, which cited “On the 
Gila, Sonora; Parry.” Kearney and Peebles (1951, p. 845) observed that the species is “ap- 
parently known in Arizona only from the type collection on the Gila River (Bigelow),” 
and the present authors have located no further records of the species from Arizona. In 
contrast, plants matching the type of C. bigelovii (below) have been collected at numer- 
ous localities in northeastern Mexico (Coahuila, Nuevo Leon, San Luis Potosi—Turner 1997) 
and at two sites in Texas (mapped in Turner et al. 2003; documented here). Eupatorium 
madrense (as cited below) from Nuevo Leon is a synonym of Eupatorium bigelovii. 
U.S.A. Texas. Brewster Co.: Big Bend National Park, Basin of Chisos Mt. base of Baldy Peak, 6 Sep 1950, Warnock 
9587 (SRSC); Big Bend National Park, Sierra Quemada, Claro 2 Spring, 29 Oct 2000, Bartel 211 (SRSC). Uvalde Co.: 
on chalk bluffs at Park Chalk Bluff along the Nueces River, ca. 15 mi NW of Uvalde, 12 Dec 1967, Correll 35433 
(LL). 
Geographic-ecological summary.—Flowering Oct-Dec. Dry limestone hills in oak wood- 
lands, talus: ca. 1OOO-1800 m. (data primarily from Texas, Coahuila, and Nuevo Leén). 
Chromolaena bigelovii (A. Gray) R. King & i Robinson, Phytologia 20:208. 1970. Eupato- 
rium bigelovit A. Gray in Torrey, Rep. | 1s. & Mex. gund. Survey, Bot. 75. 1859. Type: U.S.A. [locality uncer- 
tain but probably along the Texas-Coahuila oie 1848-1855], J.M. Bigelow 12 (probable HOLOTYPE: 
GH, photocopy!). The protologue by Gray cited “On the Gila, Sonora; Parry.” Blake (1924) and Kearney and 
Peebles (1951) noted that the ely sou be Pi aced in Arizona than nian oo labels, however, 


ona collection that i Is reasonabl i g “E. bigelovii 


y 
n. sp.” and “E. bigelovii Gray” and do not have locality information. On the original label, “Mex. Bound. 
Survey No. 12,” a note by Gray says “Bigelow—locality not recorded”; another note (B.L. Robinson’s hand- 
writing) reads that “Identical material in Paris Herb. and Berlin Herb. bears Mex. Bound. Surv, label num- 
bered (in pencil) 471.” A Parry collection that would represent a potential type for Eupatorium bigelovii 


has not been located at GI 


NE IER THE U.S.A. 1253 


The species represented by the Bigelow collection is the one that has been collected nu- 
merous times in northeastern Mexico and immediately adjacent Texas. The collection 
almost certainly was made along the Texas-Mexico boundary, in the known range of the 
species, in the region covered by the United States and Mexican Boundary Survey of 1848- 
1855, for which Bigelow worked as a naturalist. Numerous collections by Bigelow from 
southwestern Texas are specifically cited in the Survey's “Botany” report. In the tentative 
interpretation here, a mistake in manuscript preparation apparently led to a substitu- 
tion both for the correct collector (Parry instead of Bigelow) and correct locality (On the 
Gila, Sonora, instead of ‘along the Texas border’). Gray’s choice of epithet supports this 
interpretation. 

Eupatorium madrense S. Watson, Proc. Amer. Acad. Arts 26:137. 1891. Typr: Mexico. Nuevo 

Leon: Sierra Madre near Monterey, 7 Jun 1888, C.G. Pringle 2201 (HOLOTYPE: GH photocopy!). 


ACKNOWLEDGMENTS 


We are grateful to Steve Boyd and Michael Denslow (RSA-POM) for information and pho- 
tocopies of Eupatorium arborescens, Phil Jenkins and Michael Chamberland (ARIZ) for 
checking potential occurrences of Chromolaena bigelovii at ARIZ, Phil Jenkins for com- 
ments on Ageratina thyrsiflora, Jane Mygatt (UNM) and Rich Spellenberg (NMC) for pro- 
viding information on New Mexico collections, Barbara Hellenthal for checking the po- 
tential occurrence of collections at NDG, Harold Robinson (US) for review comments, 
and Emily Wood and Walter Kittredge (GH) for providing information and photocopies 
of various GH collections. 


REFERENCES 


Bake, S.F. 1924. Asteraceae. In:P.C. Standley, Trees and shrubs of Mexico.Contr, U.S. Natl. Herb.23(4):1401- 
1641. 

Grastorr, J.L. 1972.A systematic study of the North and Central American species of Stevia.Ph.D.diss., 
Univ. of Texas, Austin. 

Husearo, J.P. 1999. Penstemon pulchellus Lindl. [= P campanulatus (Cav.) Willd.]:A specious Mena or 
New Mexico's flora. New Mexico Botanist,|ssue No.11.<http://web du/~kallred/herbweb 
newpage27.htm> Accessed June 2005. 

Lene, JH. 1978. A catalogue of the flora of Arizona. Desert Botanical Garden, Phoenix, Arizona. 

McVaucu, R.1956. Edward Palmer: plant explorer of the American West. University of Oklahoma Press, 
Norman. 

McVauch, R. 1984. Compositae. Flora Novo-Galiciana 12:1-1157. 

Martin, W.C. and C.R. Hutcuins. 1980. A flora of New Mexico. Vols.1 and 2. J. Cramer, Vaduz. 

Turner, B.L. 1997. The Comps of Mexico. A systematic account of the family Asteraceae. Volume 1. 
Eupatorieae. Phytologia Mem. 11:i-iv; 1-272. 

Turner, B.L., H. NicHols, G.C. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas.Vol. 1, Dicots; 
Vol. 2, Monocots, ferns, Sida, Bot. Misc. 24. 

Wooton, E.O. 1906. Southwestern localities visited by Charles Wright. Bull. Torrey Bot. Club 33:561- 


Wooton, E.O. and PS. STanpLey. 1915. Flora of New Mexico. Contr. U.S. Natl. Herb. 19:1-794. 


1254 BRIT.ORG/SIDA 22(2) 
Book REVIEWS 
Bruct A. Bot. 2004. Hawai’i’s Native Plants. (ISBN 1-56647-066-6, pbk.). Mutual Publish- 
ing, 1215 Center Street, Suite 210, Honolulu, H1 96816, U.S.A. (Orders: 808-732-1709, 
808-734-4094 fax, mutual@mutualpublishing.com, www.mutualpublishing.com). 
$22.95, 224 pp., approx. 200 color photographs, 6" x 9" 
Dr. Bruce A. Bohm’s Hawai’i’s Native Plants presents a historical view of botany on the Hawaiian islands and is 
quite pleasant to read. I must begin by saying that Hawaii’ Native Plants is nota field guide for quick identifica 
tion of plants. Rather, it carefully introduces the geologic, ecological, evolutionary and cultural history of each 
major taxonomic group, often in too much detail for the lay reader. This book is not organized like a traditional 
field guide; species information is presented in narrative form with headers of both Genus and Hawaiian names 
introducing each group. Excellent color photographs throughout the book provide identifying characteristics of 
species, though not all species are a gaa That being said, this book would be an excellent reference / 
companion for a “quick-identification” field guide of Hawaiian plants. 


is work begins with a chapter (The islands)on the geologic, ecological and cultural history of the Hawai- 
ian islands It gives a brief introduction to each island, eens deseri ested of vegetation patterns as well as brief 
historical backgrounds. This first chapter reads almost like a travel gui Hawai'i, but is a good introduction to 
the processes and historical events that have helped shape she island 


s current flora and fauna. Bohm focuses on 
plants that are easily accessible to the amateur naturalist/tourist but does include some more challenging-to- 


lind species. His stated audience is persons traveling to Hawai’; his goal is for the reader to become appreciative 
of Hawai’'s native flora and informed of the current threats to Hawai’i’s biodiversity. However, the average tourist 
(especially if he/she has no background in biology or geology) may find this book a challenging read because of 
the detail in which Bohm discusses biological and geological processes 


ative Hawatian Plants, the second chapter, is the one that provides the taxon-specific descriptions anc 


introduction to the major groups of native Hawaiian plants; thus, it makes up the majority of the book. As men 
tioned hee ai Plants are pateedneee uInE neading> of the Genus ane raya an TAMIeIe), Plants are first de- 
scribed fori history f most groups 
are discussed. For most ai the taxa, these short literature explorations may le sad the lay reader to loose interest in 
the technical information presented (ie. phylogenetic analysis or molecular genetics techniques). 2 conservation 
status, native cultural uses, factors that have affected the colonization, distribution, and populati 
species (e.g. major storms, volcanic activity, human activity) are a great addition to this book, as this nloumation 
is rarely included to this extent in traditional field guides. The end of this chapter contains a small section en- 
titled Ferns and Fern Allies; it is short and limited to relatively easy to find species. The author suggests Daniel 
— Hawat'i’s Ferns and Fern Allies (2003) tor a more soln iE enSING guide to Sa erouD 

e third chapter, Alien Invasion, discusses the extent t I native f] | 


a Th he author includes this chapter in hopes that the reader will gain an apeeeaaen for the “full impact” 
invasive species can have on native biota. He introduces invasive plants by taxonomic group and discusses how 
each group became introduced to the islands, focusing on species considered to be invasive as opposed to natural- 
ized or introduced species. There is also memes of the ecological CHATS of invaders that have made 
es so o successful on awe iand of s that are being try to eradicate) invasives. 
final chapter (There Is No Future In Extinction) begins with a discussion af extinction in general, con- 
ete species loss and biological characteristics sist may make certain taxa more prone to extinction. In 
this chapter Bohm gives the reader an abundance of literary references on extinction for further exploration. He 


includes a brief presentation of global hotspots of plant diversity and endemism, then discusses threats specific 
to Hawaiian flora and describes specific conservation efforts to preserve or restore native populations of Hawai- 
ian plants. Finally, the author provides a short appendix at the end of the book that explains methods commonly 

sed by taxonomists to identify and study eaeee history of species, including molecular phylogenetics 
ages that have allowed for better identification of plants. 


any 


or avid natural historians with some bac nee knowledge of geology, evolution and ecology, this book 
isa Stee addition to the botanical literature on Hawai'i. Professors teaching courses on Hawaiian natural 


history or botany would find this a useful volume; it is also quite affordable for students. Hawaii's Native Plants 


would best be read in ae entirety before a trip to Bona ito eee an unde standing of the basic lationship 
between plant taxaa logy of plant groups. TI ily 


find certain species (often roadside access), and would be a usef ful tool for planning outings on trips.—J. Hayley 
Gillespie, University of Texas at Austin, Integrative Biology, Austin, TX 78712, U.S.A, hayleygillespie@mail.utexas.edu 


SIDA 22(2): 1254. 2006 


BOOK REVIEWS 1255 


MELINDA A. ZEDER, DANIAL G. BRADLEY, EvE EMSHWILLER, and Bruce D. Smitu (eds.) 2006. Docu- 
nting Domestication: New Genetic and Archaeological Paradigms. (ISBN 978-0- 
520-24638-6, hbk.). University of California Press. Berkeley, Los Angeles, London. (Orders: 
California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jer- 
sey 08618; Tel.: 609-883-1759; Fax: 609.883-7413) $70.00, 361 pp., b/w figures, b/w 
photos, tables. 
Apoues ae studies were well underway during the last century, Documenting Domestication: 
Ww haeological Paradigms definitely sets a new standard for scientific investigations, integra- 
tions, and interpretations for the 21st Century. Certainly, rapidly advancing electronic technologies for entering, 


storing, and retrieving data rapidly and accurately have ele the necessary tools to enable and facilitate 
a much larger body of assemblages and studies. More- 


research and interpretation and to utilize materials fro 
uch earns developments as high-power scanning electron microscopy and techniques “for the re- 


over, $ 
t macro- and micro-fossils” have greatly expanded and enhanced 


co identification of 
eee ae and genetic studies. 
mposium presented at the annual meetings of the American Association for the Advancement of Sci- 
ence (2001) featured four speakers, each addressing progress and future concerns and challenges in pene spe- 
cific studies “on the domestication of plants and animals from archaeological and genetic perspectives.” That 
kers are editors as well as contributors. 
omestication: Bringing Together Plants, Animals, 


provided the impetus for this volume, and three of those spea 
e overarching introductory chapter, “Documenting D 
Archaeology, and Genetics,” is written by the editors. It provides an in- de pe overview of earlier work from the 
mid 19th Century, beginning with perceptions from Darwin and through the present, with its rapidly 
expanding resources and technologies. Discussions of changing approaches and understandings of the scientific 
disciplines herald fresh new ways of ae and interpreting data, not only from newly collected evidence 


but from renewe tion sting data 
The book is designed ona ee matrix” (table 1.1). Research for domestication of plants and animals 
is documented in special sections based on archaeological studies and on Boe sas bach oe is respon- 
the genetic 


sible for one of tl gy of plant d 

documentation of plant domestication; Melinda Zeder, annie of animal domestication: and Dan Bradley, 

the genetics of plant domesticati ion 
ditor has iew chapter preced les in th egory. This is very helpful for 

the reader, and it pemmits rapid identi ication sof chapters ely 2 be relevant t to one’s own interest or research. 

1 provide informa- 


The chapters 
tion that previously has not been easily available The ue mmterest will oe be drawn first by those 


that most oe oe interest pteee and os Re me, aus I enyea the entire volun 
TI of fi | visual percep- 


tion of data and eee are Grice additions. 

ther sad lor us volume? is the wide diversity of geographic areas covered by the researchers. The range 
of d hand South America, Eurasia, and the Middle East. That breadth of research 
enhances the ee of key animal and plant coverage and is eee helpful in analyzing the data. (To date, 
most studies have been centered on are: often very small areas; coma een wath aero iaeee from eis 


sites can be very distorted. It is not easy t formulate I 


the retrieved data were based on the same or femmes procedures and markers.) 
Interestingly, the new methods, tools, data retrieval, ease o 
to new information being immediately available from remote study areas have were workers to re-think 
older, accepted interpretations and to construct solid new paradigms 
ew questions; me eae and new pecan of evidence from ancient archaeological as- 


1 | 
A data, ac 2SS to 7 inaccessible 


nd access 


semblages, « to phytoliths in plants or study 
studies. call of these things enable researchers to study op evidence again and refine or rane the interpretations. 
ning and execution of this Vohaniee are bom peas te and professional. The use of Vincent van Gogh's 


“The See for the cover is particularly p 

Scientists and historians usually concur that 
determining events in the history of mankind. This volume is 
ing.—Helen Jeude, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. 


rr L 
d tication of pants and animals is one of the major 


stand- 


SIDA 22(2): 1255. 2006 


1256 BRIT.ORG/SIDA 22(2) 


GFORGE Y ATSKIEVYCH. 2006. Steyermark’s Flora of Missouri Volume II- Revised edition. (ISBN 
1-930723-49-0, hbk.). Missouri Botanical Garden Press, P.O. Box 299, Saint Louis, MO 
63166-0299, U.S.A. (Orders: http://www bgpress.info/). $48.00, 1181 pp., b/w line 
drawings, maps, 7 3/8" X 97/8". 


fai 


yanyone interested in the Missouri F 


4 


ora Project, George Yatskievych’s revision of the original Flora of Missouri 
i Julian Steyermark was looked forward to with great anticipation. Volume 1, published in 1991, covered pteri- 
dophytes, gymnosperms, and monocots. It was so well written, organized, illustrated and thorough, that Mis- 
souri botanists anxiously awaited Volume 2. Published in June 2006, Volume 2 covers the first part of the dicots 
alphabetically from the Acanthaceae through two subfamilies of the legumes or Fabaceae (Caesalpinoideae and 
Mimosoideae). Volume 2 is so well done that the author has even surpassed the eae of Volume | and stu- 
dents of Missouri's flora have been greatly rewarded for their patience. Each species account includes a detailed 
description that is the most thorough I have ever observed in any North American floristic manual. Also pro- 
vided are blooming periods, county dot range maps, characteristic habitats, relative abundance and illustrations 
for the overwhelming majority of species. Most taxonomic treatments follow the findings outlined in most re- 
cently published literature, Sly for the most part, the rearrangement of genera within the Asteraceae that 
has been adopted in inuals (e.g., Flora of North America). 

Even without the seegone ae 2 isa monumental enue Nonet heless, the illustrations are so 
detailed and so complete that the 193 plates are the most accurate repres “treated species | have seen in 
any flora. One aspect | particularly like is the enlarged detail of the fruit illustrated ete is easy to recognize, 
especially for genera and species that are often difficult to visualize in sufficient detail to distinguish between 
closely related taxa (e.g,, species of Vernonia, plates 299 and 300; and species of Cuscuta, plates 364 and 365). 
Another highlight is that the Brassicaceae includes | ve lor both { Novwersand ED 651-659). Anyone who has 
ever tried to key out material in this family will fi ] 


a 


eS 


y helpful. Overall, the keys 
and the glossary on pages 1085-1102 covers terms tha at may be unfamiliar to some readers 
me 2 is such a quality compilation of information that it is hard to find fault. Nonetheless, Gee area 
few minor points that should be brought to light. The most obvious oversight is that the numerous genera and 
species within the Asteraceae are arranged by Tribe. Although this alignment is helpful in simplifying keying 
exercises (pages 167-171), it is extremely frustrating for readers who do not know what tribes various genera are 
ascribed, especially given the recent taxonomic changes within the oles To locate a eae genus within the 
Asteraceae, the reader must locate it by thumbing through 42 pages of index (pages | 139-1181). A potential solu- 
tion would have been to list all genera within each tribe near the - ginning of the discussion on the Asteraceae. 
Another shortcoming of the book is that the treatment does not cover the last subfamily of the Fabaceae (ie., 
Faboideace ) which includes most of the species of legumes documented for a state, Obvionsy ay writer of a 
} | } 


book this size establish a cut off somewhere, but it would have | family to b 


function wel 


covered together Alsou 1 undoubtedly a selfish and unrealistic expectation of his reviewer, it would be lave 
been helpful to include a key that included all dicot families, especially given the length of time that is likely 
required to complete the last treatment. With any book of such magnitude, minor errors are sure to be inherent 
but only a few have been identified to date. The couplets separating Helenium autumnale from H. virginicum in 
the key on page 490 were inadvertently transposed. Helenium virginicum should be 3a, not 3b. On plate 386 
(page 1067), the fruiting branch illustrated for Senna obtusifolia should be labeled 10e and the fruiting branch for 
S.occidentalis should be labeled as 10f. 


In conclusion, Ste yermark’s Flora of Missouri- Volume 2 is nothing less than a botanical masterpiece in print 

and it clearly warrants a spot on the shelf of every botanist in Missouri. Because its application and coverage in- 

ludes many species that have been documented for many states in the Midwest and eastern Great Plains, it would 

make an excellent reference book for anyone studying plants in the eastern United States.—Paul M. McKenzie, 

U.S. Fish and Wildlife Service, 101 Park DeVille Dr; Suite A, Columbia, Missouri 65203, U.S.A., Email: 
paul_ mckenzie@fwsygov 


— 


SIDA 22(2): 1256. 2006 


REPORT AND UPDATE 2006 


SIDA, CONTRIBUTIONS TO BOTANY 
Source of current research in classical and modern 
botany for readers throughout the world. 


systematic 
...bringing out the best in botanical science for plant conservation and education 

To all authors, we thank you for choosing Sida, Contributions to Botany as your forum to 
disseminate information and knowledge gained by scientific inquiry. Volume 22, com- 
plete in two numbers, will be the last volume of the journal title. Volume 22 has 1274 
pages, 169 authors, 105 published contributions, and includes 97 new names and new 
combinations. Published papers are available online in Adobe Acrobat format (PDF files); 
the PDF files are true representations of the hard copy of Sipa <http:// www.britorg/sida/ 
SCBCurlssue.htm>. 

A word of thanks to the 298 individuals acknowledged on pages 1258-1259, who 
generously supported Sip through their time and expertise in reviewing 226 manuscripts 
submitted for volume 22. All manuscript submissions are peer-reviewed by distinguished 
reviewers; your support is deeply appreciated. If by chance you reviewed a manuscript 
and your name was left out, the error rests solely on the shoulders of this editor and | 
truly apologize. Please bring any omissions to my attention. 

Below are the dates of publication for the two issues of volume 22, subscriptions. We 
also bring to your attention (see bottom of this page) the Mary M. Hennen Scientific Pub- 
lications Endowment, established to provide support to the future of BRIT’s scientific publi- 
cation program (Sida, Contributions to Botany and Sida, Botanical Miscellany). The index 
to volume 22 (2006) follows. Guidelines to contributors are available online at http:// 
www.britorg/sida/SubmitPaperhtm. 

We thank all authors, reviewers, subscribers (individuals, institutions, organizations), 
and readers for your continued interest and support. It is our plan to continue bringing 
you the best sources of current research in classical and modern systematics with your 
continued support. Wishing you the best in 2006 and beyond.—Barney Lipscomb (BRIT), 
Editor: John W. Thierett (NKU), Associate Editor; and Félix Llamas (LEB), Contributing 
Spanish Editor. 


Dates of publication 
22(1), pp. 1-846: 11 August 2006 
22(2), pp. 847-1274: 23 November 2006 
Mary M. Hennen Scientific Publications Endowment Established.—In honor of his wile, 
Mary, BRIT research associate, professor emeritus of botany at Purdue University, the 
world’s pre-eminent authority on the systematics and biogeography of the rust fungi, Joe 
Hennen has established the Mary M. Hennen Scientific Publications Endowment to pro- 
vide support to the future of BRIT’s scientific publication program. Joe and Mary (life- 
science librarian at Purdue University) have long recognized that publication of scien- 
tific botanical information is an integral part of conserving our natural heritage. 

Every gift to the endowment will make a difference for the future of BRIT scientific 
publications. There are many different ways to make a gift; use a credit card, give a check, 
stock, pledge, bequest, trust, memorials, employer/employee matching contributions, 


SIDA 22(2): 1257. 2006 


1258 BRIT.ORG/SIDA 22(2) 


installments, etc. The BRIT Press seeks your support to ensure innovation and excellence 
in preparation, manufacture, and distribution of botanical research and scientific dis- 
coveries for the twenty-first century. For more information about the BRIT Press or any- 
one interested in making a contribution to the Mary M. Hennen Scientific Publications 
Endowment—which supports Sida and Sida, Bot. Misc.—contact Barney Lipscomb at the 
Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, 
US.A., 817-332-7432; barney @brit org; sida@britorg. 


REVIEWERS: VOLUME 22 (2006) 
298 reviewers: several individuals reviewed more than one manuscript 
Thank you for supporting Sida, Contributions to Botany, 


Acedo, Carmen 


Allred, Kel ly W. 
Anderson, Loran C. 
Andreas, Barbara K. 
Arnold, A. E 
Arriagada, Jorge E. 
Austin, Daniel F 
Ayala-Silva, Tomas 


— 


izabeth 


Ayers, Tina J. 
Bailey, Jr, Claude J. 
Baird, Gary L. 
Baldwin, Bruce G. 
Ball, Peter W. 
Ballard, Jr, Harvey E. 
Ballenger, Julie 
Barber, Susan C. 
Barkworth, Mary E. 
Barlow-lrick, Patricia 
Barrie, Fred R 
Baskin, Carol C. 


Bz 
S 
> 
or) 


Brown, Larry E. 
Brunell, Mark S. 
Buck, William R 
Burnham, Robyn 
Burns, Jean 


SIDA 22(2): 1258. 2006 


Campbell, Julian 
Cardellina, John 
Carlquist, Sherwin 
Carlsward, Barbara S. 
Carranza, Eleazar 
Carstens, Kenneth C. 
Carter, Richard 


Chambers, Kenton L. 
Chinnappa, C 
Christensen, a Ib 
Church, Sheri A 
Clark, Lynn G. 
Columbus, J. Travis 
Consaul, Laurie | 


Cruden, Robert W. 
Davidse, Gerrit 
Davila, Patricia D. 
Delprete, Piero G. 
Denslow, Michael W. 
Dorr, Laurence J, 
Dowling, R.M. 
Duvall, Jams 

Eggli, Urs 

Elisens, Wayne J. 
Endress, Mary E. 
Essig, Frederick B. 
Evans, Dan K 
Ezcurra, Cecilia 
Fantz, Paul R. 
Farmer, Susan B. 
Ferguson, C -aroly nj. 
Fernandez-Concha, Ger- 
man Carnevali 

Fishbein, Mark 

Ford, Bruce A. 


Ford-Werntz, Donna 
Freeman, Crai 

Freire, Susana Edith 
Freire-Fierro, Alina 
Frye, Christopher 
Fryxell, Paul A. 
Galloway, Cynthia M. 
Gandhi, Kanchi N. 
George, Robert 


German Ramirez, Maria T. 


Ghatoor, Abdul 


Gomez-Sanchez, Maricela 
Gonzalez-Elizondo, M. 
Socorro 

Gralt, Pamela 

Grant, Verne 

Hall, David W. 

Hallgren, Steven 

Hansen, Bruce F 

Hat e 
en Reuse L. 


> 


Hatch, Stephan L. 
Haynes, Robert R. 
Hennen, Joe 
Henrickson, James 
Herrera Arrieta, Yolanda 
Hess, Bill 
Hill, Steven R. 
Hoch, Pete 
Hoggard, Gloria 
Hoggard, Ronald K. 
Holmes, Walter C. 
Holst, Bruce K. 
Horne, Francis R. 
Howcrott, N.HS. 
Humber, Richard A. 
Hyatt, Philip 


Janovec, John 
Jarvis, Charlie 
Johns, Bob 
Johnson, Dale E. 
Johnson, George P. 
Jones, Thomas H. 
Judd, Walter S. 


Kaul, Robert B. 
Kawasaki, Lucia 
comes Sterling C. 


Kennedy, Robin C. 
Kephart, Susan R. 
Khan, Zaheer 
Kirkbride, Jr, Joseph H. 
Knapp, Sandra 

Kral, Robert 
Krapovickas, Antonio 
Kumar, Muktesh 
Kumar, Muktesh 


Lammers, Thomas G. 
t, Eric 


Landrum, Leslie R. 
Lee, Linda 
Lemke, David E. 


Liogier, Main 
Lipkin, Robert 
Liston, Aaron 
Livshultz, Tatyana 
Llamas, Félix 
Lohmann, Lucia G. 
Lonard, Robert I. 
Lowry IL, Porter P. 
MacDougal, John 
Macklin, James A. 


REVIEWERS 


Magill, Robert E. 
Magrath, Lawrence K. 

Malmquis 

oe le 

Washington 

Martin, David L 

Martinz Diaz, Mahinda 


M Teresa 
Mellichamp, T. Lawrence 
Middelton, David 
Mill, Robert 

Molpus, Jane 

Moore, Gerry 
Morefield, James D. 
Moreno, Nancy P. 
Mueller, Jochen 
Murray, David F. 
Murrell, Zack 

Naczi, Robert EC. 
Nasir, syed Mahmood 


Ne : 
Nepokroeff, Molly 
Nesom, Guy 
Nevling, Lorin L 
Nicolson, Dan H. 
Nordman, Carl 
Norris, Dan 


SIDA 22(2): 1259. 2006 


Noyes, Richard D. 
Oberlies, Nicholas H. 
Olmstead, Richard G. 
Orzell, Steve L 
Palmer, Michael W. 
Paratley, Robert 

Peck. lal 


pceom, Paul M 


poner ie Kristen E. 


Powell, A. Michael 
Pratt, Donald B. 
Pringle, James 5. 
Pruski, John F 
Pyne, Milo 

Qiu, Yin-Long 
Rabeler, Richard 


Robertson, Kenneth R. 


Rock, Janet H. 
Rohrer, Joseph R. 
Rossman, Amy 


Rumely, John I. 
Rutishauser, R. 
Ryburn, Adam K. 

Saar, Dayle E. 

Salazar, Jackeline 
Sanchez - Pino, Ivonne 
Sander 

ae var E 
Schultz, Joanna 

Schuyler, aed aes 
Semple, ee 


Shaw, Joe 

Sheviak, les 
Shulkina, Tatyana 
Siddiqi, Akma 
Simpson, Beryl B. 


Smith, Latimore 
Smith, Marian 
Smith, S. Galen 
Soltis, Douglas E. 


Steury, Mr. Brent 
Stevens, G.N. 
Stewart, Scott 


Syring, John 


1259 


Taylor, Charlotte M 
Taylor, David D. 
Telfair IT, Ray C. 
Thiede, Joachim 


Umber, Ray 


Van Horn, Gene S. 
Vander pool Staria Scott 


Wagner, Warren L. 
Wallnofer, Bruno 
Ware, Stewart 
Weakley, Alan S. 
Webster, Grad 
Weigend, Maximilian 
Wilken, Dieter H. 
Wilkie, Peter 
Williams, Justin 
Wilson, Barbara L. 
Wilson, Carol A. 
Wipff, Joseph K. 
Witsell, Theo 
Wunderlin, Richard P. 


Yang, TY. Aleck 
Yatskievych, George A. 
Yokota, Masatsugu 


udio, Sergio 
Zarucchi, James L. 
Zmarzty, S 
Zona, Scott A. 


1260 


BRIT.ORG/SIDA 22(2) 


INDEX TO 103 TITLES WITH 148 AuTHoRS: VOLUME 22 (2006) 


We know you have a choice. Thank you for choosing and 
supporting Sida, Contributions to Botany. 


A demographic and ecological analysis of 
Geocarpon minimum (Caryophyllaceae): a 
federally threatened species in southwest Mis- 
souri by Tim SmitH and JoserH S. ELy—22(2):1145 

A new combination in Polytrichastrum 
(Bryophyta: Polytrichaceae) by G.L. Smit Mer- 
Ri —22(1):547 

A new combination in Tetraphis (Bryophyta: 
Tetraphidaceae) by JupitH A. Harpe. —22(1): 
549 

A new combination in the Bromus catharticus 
complex (Poaceae: Bromeae sect. Cerato- 
chloa) by Ana Maria PLANCHUELO—22(1):555 

A new hybrid genus and 11 new combina- 
tions in North American grasses by Mary E. 
BARKWORTH—22(1):495 

A new name for Sida, Contributions to Botany 
(1962-2006) by Barney Liescomsa—22(2):847 

A new narrowly endemic species of Clematis 
(Ranunculaceae: subgenus Viorna) from 
northeastern Texas by Dwayne Estes —22(1):65 

Anew species of Boltonia (Asteraceae) from the 
Ridge and Valley physiographic province, 
U.S.A. by JOHN F. Townsenp and Vesna KARAMAN- 
CASTR —22(2):873 

A new species of Dodecatheon (Primulaceae) 
from the northern coast range of Oregon and 
Washington by Kenton L. CHAmBeERS—22(1):461 

A new species of Eugenia (Myrtaceae) from the 
Monteverde region, Costa Rica by Frep R. 
Barrit-—22(2):1071 

Anew species of Pediomelum (Fabaceae) from 
the lower Piedmont Plateau of Georgia and 
South Carolina by James R. ALUSON, MICHAEL WAYNE 
Morris, and AsHtey N. EcAN—22(1):227 

Anew species of Stenanthium (Melanthiaceae) 
from Tennessee, U.S.A. by B. Eucene Worrorp— 
22(1):447 

Anew species of waterlily (Nymphaea minuta: 
Nymphaeaceae) from Madagascar by KennetTH 
LANDON, RicHarD A. Eowarps, and P. Ivan Nozaic— 
22(2):887 


SIDA 22(2): 1260. 2006 


A new species of witch-hazel (Hamamelis: 
Hamamelidaceae) apparently endemic to 
southern Mississippi by S.W. Leonarpb— 
22(2):849 

A quantitative study of styles and achenes of 
terminal and basal flowers — of 
Schoenoplectus hallii (Cyperaceae), a rare 
plant species of transient wetland habitats by 
Marian SMITH, SARA AMMANN, Nancy Parker, and 
Paice Meriier-CHERRY—22(2):1159 

A vascular plant inventory of Starkey Wilderness 
Preserve, Pasco County, Florida by Emity Fercu- 
SON and RicHarb P. WuNDERLIN—-22(1):635 

Amelichloa: a new genus in the Stipeae 
(Poaceae) by Mirta O. ArriaAca and Mary E. 
BaRKWORTH—22(1):145 

An updated, annotated vascular flora of Caddo 
Parish, Louisiana, with notes on regional phy- 
togeography and ecology by Barsara R. 
MacRoserts and MicHaet H. MaAcRoserts 
—22(1):119] 

Annotated checklist of the vascular flora of the 
Big Sandy Creek Unit, Big Thicket National Pre- 
serve, Texas by Larry E. Brown, Barsara R 
MacRoserts, MicHAEL H. MacRoserts, PAUL A. 
Harcomee, Warren W. Pruess, |. SANDRA Etsik, and 
STANLEY D. JONES —22(1):705 

Annotated checklist of the vascular flora of the 
Lance Rosier Unit, of the Big Thicket National 
Preserve, Hardin County, Texas by Larry E. 
Brown, BarsarA R. MacRoserts, MICHAEL H. 
MacRoserts, Paut A. Harcomee, Warren W.Pruess, |. 
SANDRA Etsik, and Stanvey D. Jones —22(2):1175 


Big-fruited buckthorn, Sideroxylon macro- 
carpum (Sapotaceae),a long-forgotten Geor- 
gia endemic by James R. ALuison—22(1):243 

Bromus ayacuchensis (Poaceae: Pooideae: 
Bromeae),a new species from Peru, with a key 
to Bromus in Peru by Jerrery M. Saareta, Paul 
M. Peterson, and Nancy F, Rerutio-Ropricuez— 
22(2):915 


INDEX 


Capraria mexicana (Scrophularia¢ eae) in Cam- 
eron County, Texas: rediscovered in the United 
States by AtFred RICHARDSON and Ken Kinc— 
22(2):1235 

Carex bicknellii (Cyperaceae) new to Arkansas 
by Pau M. McKenzie, C. THeo WirseLt, and Joe 
Woo eriGHT—22(1):801 

Carex reznicekii, a new widespread species of 
Carex section Acrocystis (Cyperaceae) from 
eastern North America by Davin A. WerieR— 
22(2):1049 

Carex stipata (Cyperaceae), Chamaesyce 
gereyi (Euphorbiaceae), Eurytaenia texana 
(Apiaceae), Pediomelum esculentum 
(Fabaceae), and Talinum calycinum 
(Portulacaceae) deleted from the Louisiana 
flora by Barsara R. MacRoserts and Michael H. 
MacRoserts—22(2):1 221 

Cayratia japonica (Vitaceae) new to North 
Carolina and an updated key to the genera of 
Vitaceae in the Carolinas by ALEXANDER KriNcs 
and Roseet J. RICHARDSON—22(1):813 

Chromosome numbers for western and arctic 
North American species of Antennaria 
(Asteraceae: Gnaphalieae) by Jerry G 
CHMIELEWSKI—2.2(1):561 

Clematis morefieldii (Ranunculaceae) new to 
Tennessee by Dwayne Estes and Curls FLeminc— 
22(1):821 

Comments on the Gerbera-complex 
(Asteraceae: Mutisieae) by Hans V. HANsen— 
22(1):539 

Crataegus series Bracteatae and Triflorae (Ro- 
saceae) by J.B. PHipes, R.LaNce, and K.A. Dvorsky 
22(2):1009 

Crataegus series Parvifoliae and its putative 
hybrids in the southeastern United States by 
J.B. Puipes and K. Dvorsky-—22(1):423 


Dendrophorbium restingae (Asteraceae: 
Senecioneae), a new species from Sao Paulo, 
Brazil by Aristonio M. Tees, Jimi N. NakAJima, and 
Joao R. STEHMANN—22(1):123 

Drepanostachyum falcatum var. seng- 
teeanum: identity and origins (Poaceae: 


SIDA 22(2): 1261. 2006 


1261 


Bambusoideae) by C.M.A. STAPLETON— 
22(2):1081 


Ericameria nauseosa subsp. ammophila 
(Asteraceae), a new rabbitbrush from the San 
Luis Valley of Colorado by Loran C. ANDERSON 
22(2):866 

Eriogonum callistum (Polygonaceae), a new 
species from the Tehachapi Mountains of Cali- 
fornia by James L. ReveAaL —22(2):857 

Erodium malacoides (Geraniaceae), new to 
Texas by Davio E. Lemke and Jeo L. APLAcA— 
22(2):1 245 

Estudios en las Apocynaceae neotropicales XXIV: 
Bahiella (Apocynoideae: Echiteae), un 
desapercibido nuevo género endémico de 
Bahia, Brasil by J. Francisco Morates—22(1):333 

Estudios en las Apocynaceae neotropicales 
XXV: novedades y nuevos reportes en las 
Apocynaceae (Apocynoideae: Rauvolfioideae) 
de Venezuela by J. Francisco MorAtes— 
22(1):355 

Evidence for hybridization between two sympa- 
tric violet species, Viola grahamii and V. 
hookeriana (Violaceae), in central Mexico by 
Aurea C. Cortts-Patomec and Harvey E. BALLARD, 
Jp,—22(2):1119 

Exotic plant introduction in Kansas, two new 
species by Iracee BARNARO—22(1):777 


Floristic diversity and composition of terra firme 
and seasonally inundated palm swamp for- 
ests in the Palma Real Watershed in lower 
Madre de Dios, Peru by FerNANbo H. CorNeJO 
VALVERDE, JOHN P JANovec, and MatHias W. TosLer— 
22(1):615 

Four novelties and a lectotypification in Matelea 


(Apocynaceae: Asclepiadoideae) from His- 
paniola by ALexaNDeR KRINGS—22(2):941 

Fungi associated with Borrichia frutescens 
(Asteraceae): insect galls and endophytes by 
Diane Te StRAKE, Amy Hapbock Keacy, and Peter D. 
STILING = 22(1):755 

Galium tricornutum (Rubiaceae) and Parentu- 
cellia viscosa (Scrophulariaceae) new to Okla- 
homa by Amy K.ButHoo and Bruce W. HoaclAND— 
22(2):1235 


1262 


Gratiola brevifolia (Plantaginaceae) new to the 
flora of Delaware, the Delmarva Peninsula, and 
the Mid-Atlantic by Westey M. Knapp and Dwayni 
Estes —22(1):825 

Green fringed orchid (Platanthera lacera, 
Orchidaceae) in southern Louisiana by Cuaries 
ALLEN, SARA THAMES, HOWARD ANDERSON III, BILL NEw- 
FON II], RHONDA Hampton, and Georce FisHeR— 
22(1):805 


Hedyotis australis (Rubiaceae) new to Missouri 
and Florida and related species in the south- 
central United States by Watter H. Lewis— 
22(1):831 

Herbivory of feral goats on Espiritu Santo Island, 
Gulf of California, Mexico by Jose Luis Leon-pe 
LA Luz and Reymunno Domincuez-CADENA— 
22(2):1135 

High levels of seed inviability among seven 
populations of the endangered Short's gold- 
enrod (Solidago shortii: Asteraceae) by 
Patrick J. Catie, Ropert F.C. Naczi, CHRISTINA 
SHACKLEFORD, aNd Jessica CaiceDO—22(1):735 

Hill cane (Arundinaria appalachiana), a new 
species of bamboo (Poaceae: Bambusoideae) 
from the southern Appalachian Mountains by 
J.K. Tripcett, A.S. Weaxtey, and L.G. Ciark— 
22(1):79 

John W. Thieret (1926-2005) by Rateu L. 
THOMPSON—2.2(1):3 

John W.Thieret, a student's perspective (1967- 
2005) by Rosert R. HAyNeS—22(1):25 

John W. Thieret, associate editor of Sida (1972- 
2005) by Barney Lipscomp—22(1):55 

John W. Thieret, colleague and editor friend 
(1969-2005) by Wittiam F.MaHter—22(1):51 

John W. Thieret, Dedication—22(1):1 

John W.Thieret, the curator by Rosert F.C.Naczi— 
22(1):21 

John W.Thieret, valuable botanical friend (1965— 
2005) by Ronato L. Stuckey —22(1):33 

More American black sapotes: new Diospyros 
(Ebenaceae) for Mexico and Central America 
by Mircuett C. Provance and Anorew C. SANDERS— 
22(1):279 


SIDA 22(2): 1262. 2006 


BRIT.ORG/SIDA 22(2) 


Morphometric analysis of an Amelanchier (Ro- 
saceae: Maloideae) complex on the Delmarva 
Peninsula (Delaware, Maryland and eastern 
Virginia) resolves the taxonomic identities of 
Amelanchier obovalis and A. canadensis by 
CHRISTOPHER T. Frye —2.2(2):1027 


Naturalization and extirpation of water hyacinth 
(Eichhornia crassipes, Pontederiaceae) in 
southwestern Arkansas, U.S.A. by RENN TUMLISON 
and Brett Serviss—22(1):749 

New and disjunct records of Eleocharis liesneri 
(Cyperaceae) from South America by Davio J. 
RoseN—22(1):781 

New and noteworthy collections for Arkansas by 
James H. Peck and Brett E. Serviss —22(1):817 

New and significant records of vascular plants 
for Florida and for Collier County and Lee 
County, Florida by Grorae J. Witber and MartHa 
R. McComas—22(1):787 

New combinations in the genus Cymopterus 
(Apiaceae) of the southwestern United States 
by Ronato L. HarTMAN—22(2):955 

New distribution records for Eupatorieae 
(Asteraceae) in the United States by Guy L. 
Nesom and Bittie L. Turner —22(2):1 249 

New species of Myrtaceae from Ecuador and 
Peru by Bruce K. Hoist and Maria LUCIA KAWASAKI— 
22(2):931 

New taxa and combinations in cultivated bam- 
boos (Poaceae: Bambusoideae) by C.M.A. 
STAPLETON—2.2(1):331 

New vascular plant records for New Mexico by 
RONALD L. HARTMAN, BriAN Rel, B.E. Nevson, and BriAN 
JACOBS—=2.2(2):1225 

Notes on southwestern Moraceae by ALAN T. 
WHITTEMORE—2.2(1):769 

Notes on the life and work of James Brigham 
McFarlin, Florida botanist by Tom Patmer— 
22(1):608 

Notes on types in Apocynaceae: Asclepia- 
doideae in Cuban herbaria and four lectotypi- 
fications in West Indian Gonolo-binae by 
ALEXANDER KRINGS and Paut R. Fantz—22(1):533 

Novedades en Gentianaceae para América del 


INDEX 


Sur by Eva M. FivippA and Gtoria E. BARBOZA— 
22(1):129 

Poa matri-occidentalis (Poaceae: Pooideae: 
Poeae:Poinae),a new species from Mexico by 
Pau M. Peterson, Rosert J. SorENG, aNd YOLANDA 
Herrera ARRIETA —22(2):904 

Psidium cauliflorum (Myrtaceae), a new spe- 
cies from Bahia, Brazil by Leste R. LanbruM and 
Marcos SopraL—22(2):927 


Quadruple, triple, double, and simple pappi in 
the goldenasters, subtribe Chrysopsidinae 
(Asteraceae: Astereae) by JoHn C. SemPLe— 
22(1):503 


Range expansion of Hexalectris grandiflora 
(Orchidaceae) in Texas by MarcaretT BROWN- 
Marsden and Anne B. CoLLins—22(2):1239 

Rejection of lectotypification of Aster amellus 
(Asteraceae: Astereae) and selection of a new 
lectotype by JoHN C. Semele —22(2):1087 

Review of Crataegus series Pulcherrimae (Ro- 
saceae) by J.B. PHipes, RJ. O’Kennon, and KA. 
Dvorsky —22(2):973 

Revision of Bauhinia subgenus Bauhinia sec- 
tion Amaria (Cercideae: Caesalpinioideae: 
Fabaceae) by RicHarp P. WuNDERLIN—22(1):97 

Root-shoot anatomy and post-harvest vegeta- 
tive clonal development in Lophophora 
williamsii (Cactaceae: Cacteae): implica- 
tions for conservation by Martin Terry and JAMES 
D. MausetH—22(1):565 


Sexual and apomictic prairie fleabane (Erigeron 
strigosus) in Texas: geographic analysis and 
a new combination (Erigeron strigosus var. 
traversii, Asteraceae) by RicHarb D. Noyes, Helo! 
GERLING, and CarLA VANDERVOORT—22(1):265 

Seymeria falcata (Scrophulariaceae), a new 
record for Texas and the United States by 
JOSELYN FENSTERMACHER——22(1):8 11 

Six new combinations in Dodecatheon 
(Primulaceae) by James L. ReveAt —22(2):863 

Six new species of Sphagnum (Bryophyta: 
Sphagnaceae) from North America by RicHarD 
E. ANoRUS—2.2(2):959 


SIDA 22(2): 1263. 2006 


1263 


Taxonomic overview of the Heterotheca villosa 
complex (Asteraceae: Astereae) by Guy L. 
Nesomg—22(1):367 

Taxonomic review of Symphyotrichum patens 
(Asteraceae: Astereae) by Guy L. Nesom— 
22(2):1075 

Taxonomy and conservation of medicinal plants 
in canal-irrigated areas of Punjab, Pakistan by 
KHALID Faroog Aksar and MoHAMMAD ATHAR— 
22(1):593 

Taxonomy of Cuscuta gronovii and Cuscuta 
umbrosa (Convolvulaceae) by MiHAl Coste, 
Guy L. Nesom, and Sasa STEFANOVIC—22(1):197 

Taxonomy of Lantana sect. Lantana 
(Verbenaceae): |. correct application of Lan- 
tana camara and associated names by Roctr 
W. SANDERS —22(1):381 

Taxonomy of North American species of 
Oldenlandia (Rubiaceae) by Eowarn E. Terret 
and Harotp Rosinson—22(1):305 

Taxonomy of the Cuscuta indecora (Convo- 
lvulaceae) complex in North America by 
Munat Coste, Guy L.Nesom, and Sasa STEFANOVIC— 
22(1):209 

Taxonomy of the Cuscuta pentagona complex 
(Convolvulaceae) in North America by Mina\ 
Costea, Guy L. Nesom, and Sasa STEFANOVIC— 
22(1):151 

Taxonomy of the Cuscuta salina-californica 
complex (Convolvulaceae) by Mixai Costes, Guy 
L. Nesom, and Sasa SterANOVIC—22(1):177 

The lectotypification and 19" century history of 
Croton alabamensis (Euphorbiaceae s.s.) by 
KENNETH J. WurDACK—2.2(1):469 

The north-south transition of flora across Arkan- 
sas:a preliminary phytogeographical analysis 
by MicHaeL H. MacRoserts and Bargara R. 
MacRoserts—22(1):725 

Thomas Walter Typification Project, |. Observa- 
tions on the John Fraser folio by Daniet B. 
Ward—22(2):1111 

Trichome morphology in selected Mexican red 
oak species (Quercus section Lobatae) by M. 
Lucia VAZQUEZ —22(2):1091 

Two new combinations in Peyritschia (Poaceae: 


wa 


1264 
Pooideae: Aveninae) by Victor L. Finot, Paut M. 
PETERSON, aNd FERNANDO O. ZULOAGA—2.2(2):895 


Two new species of Elymus (Poaceae) in the 
southern U.S.A. and other notes on North 
American Elymus species by Jutian J.N. 
CamPBetL—22(1):485 

Type localities of vascular plants first described 
from Ohio: supplement by James S. Princie— 
22(1):765 


BRIT.ORG/SIDA 22(2) 


Validation of the name Dahlia sublignosa 
(Asteraceae) by Dayie E. Saar and Paut D. 
SORENSEN=2.2(1):545 

Vascular floras of Sonoita Creek State Natural 
Area and San Rafael State Park: Arizona’s first 
natural-area parks by Steven P. McLauGHLiN— 
22(1):661 

Zeuxine pantlingjii, sp. nov. (Orchidaceae), a 
new species from India by Avistiek BHATTACHARIEE 
and H.J. CHOWDHERY—22(2):935 


INDEX OF 148 AUTHORS: VOLUME 22 (2006) 


Thank you for choosing Sida, Contributions to Botany. 


Akbar, Khalid Farooq—22(1):593 
Allen, Charles —22(1):805 
Allison, James R.—22(1):227, 243 
Ammann, Sara—22(2):1 159 
Anderson Ill, Howard—22(1):805 
Anderson, Loran C.—22(2):866 
Andrus, Richard E.—22(2):959 
Aplaca, Jed L—22(2):1 245 
Arriaga, Mirta O.—22(1):145 
22(1):593 


Athar \Mohammad 


Ballard, Jr, Harvey E.—22(2):1119 
Barboza, Gloria E—22(1):129 
Barkworth, Mary E.—22(1):145, 495 
Barnard, lralee—22(1):777 

Barrie, Fred R.—22(2):1071 
Bhattacharjee, Avishek—22(2):935 
Brown, Larry E—22(1):705; 22(2):1175 
Brown-Marsden, Margaret—22(2):1239 
Buthod, Amy K.—22(2):1235 


Caicedo, Jessica—22(1):735 

Calie, Patrick J—22(1):735 

Campbell, Julian J.N.—22(1):485 
Chambers, Kenton L.—22(1):461 
Chmielewski, Jerry G.—22(1):561 
Chowdhery, H.J—22(2):935 

Clark, L.G—22(1):79 

Collins, Anne B.—22(2):1239 

Cornejo Valverde, Fernando H.—22(1):615 
Cortés-Palomec, Aurea C—22(2):1119 
Costea, Mihai—22(1):151, 177, 197, 209 


sees 


SIDA 22(2): 1264. 2006 


Dominguez-Cadena, Reymund 22(2): 1135 
Dvorsky, KA.—22(1):423; 22(2):973, 1009 


Edwards, Richard A—22(2):887 
Egan, Ashley N.—22(1):227 

Elsik, |. Sandra—22(1):705; 22(2):1175 
Ely, Joseph S$ —22(2):1145 

Estes, Dwayne—22(1):65, 821,825 


Fantz, Paul R—22(1):533 
Fenstermacher, Joselyn—22(1):811 
Ferguson, Emily—22(1):635 

Filippa, Eva M.—22(1):129 

Finot, Victor L—22(2):895 

Fisher, George—22(1):805 

Fleming, Chris—22(1):821 

Frye, Christopher T.—22(2):1027 
Gerling, Heidi —22(1):265 

Hampton, Rhonda—22(1):805 
Hansen, Hans V.—22(1):539 
Harcombe, Paul A.—22(1):705;22(2):1175 
Harpel, Judith A—22(1):549 
Hartman, Ronald L.—22(2):955, 1225 
Haynes, Robert R—22(1):25 

Herrera Arrieta, Yolanda—22(2):904 
Hoagland, Bruce W.—22(2):1 235 
Holst, Bruce K.—22(2):931 


Jacobs, Brian—22(2):1225 
Janovec, John P-—22(1):615 
Jones, Stanley D.—22(1):705; 22(2):1175 


2873 


— 


Karaman-Castro, Vesna—22(2 


INDEX 


Kawasaki, Maria Lucia—22(2):931 

Keagy, Amy Haddock—22(1):755 

King, Ken—22(2):1239 

Knapp, Wesley M.—22(1):825 

Krings, Alexander—22(1):533, 813; 22(2):941 

Lance, R.—22(2):1009 

Landon, Kenneth—22(2):887 

Landrum, Leslie R—22(2):927 

Lemke, David E.—22(2):1245 

Leonard, S.W.—22(2):849 

Lewis, Walter H—22(1):831 

Lipscomb, Barney—22(1):55; 22(2):847 

Luis Leén-de la Luz, José—22(2):1135 

MacRoberts, Barbara R.—22(1):/05, 725; 
22(2):1175, 1191, 1221 

MacRoberts, Michael H.—22(1):705, 725; 
22(2):1175,1191,1221 

Mahler, William F—22(1):51 

Mauseth, James D.—22(1):565 

McCombs, Martha R.—22(1):787 

McKenzie, Paul M.—22(1):801 

McLaughlin, Steven P-—22(1):661 

Merrill, L. Smith—22(1):547 

Mettler-Cherry, Paige—22(2):1159 

Morales, J. Francisco—22(1):333, 355 

Morris, Michael Wayne —22(1):227 

Naczi, Robert F.C.—22(1):21, 735 

Nakajima, Jimi N.—22(1):123 

Nelson, B.E—22(2):1225 

Nesom, Guy L.—22(1):151, 177, 197, 209, 367; 
22(2):1075, 1249 

Newton Ill, Bill —22(1):805 

Noyes, Richard D.—22(1):265 

Nozaic, P. lvan—22(2):887 


O’'Kennon, R.J.—22(2):973 


Palmer, Tom—22(1):608 

Parker, Nancy—22(2):1 159 

Peck, James H—22(1):817 

Peterson, Paul M.—22(2):895, 904,915 
Phipps, J.B. —22(1):423; 22(2):973, 1009 
Planchuela, Ana Maria—22(1):555 
Pringle, James S—22(1):765 


SIDA 22(2): 1265. 2006 


1265 


Provance, Mitchell C.—22(1):279 
Pruess, Warren W.—22(1):705;22(2):1175 


Refulio-Rodriguez, Nancy F—22(2):915 
Reif, Brian —22(2):1225 

Reveal, James L.—22(2):857, 863 
Richardson, Alfred —22(2):1 239 
Richardson, Robert J.—22(1):813 
Robinson, Harold —22(1):305 

Rosen, David J.—22(1):781 


Saar, Dayle E—22(1):545 

Saarela, Jeffery M—22(2):915 

Sanders, Andrew C.—22(1):279 
Sanders, Roger W.—22(1):381 

Semple, Jonn C.—22(1):503; 22(2):1087 
Serviss, Brett E-—22(1):749, 817 
Shackleford, Christina—22(1):735 
Smith, Marian—22(2):1 159 

Smith, Tim—22(2):1145 

Sobral, Marcos—22(2):927 

Soreng, Robert J.—22(2):904 

Se@rensen, Paul D.—22(1):545 

Stapleton, C.M.A.—22(1):331;22(2):1081 
STEFANOVIC, SASA—2.2(1):151, 177, 197, 209 
Stehmann, Jodo R.—22(1):1 23 

Stiling, Peter D—22(1):755 

Stuckey, Ronald L—22(1):33 


Te Strake, Diane —22(1):755 
Teles, AristOnio M.—22(1):123 
Terrell, Edward E.—22(1):305 
Terry, Martin—22(1):565 
Thames, Sara—22(1):805 
Thompson, Ralph L—22(1):3 
Tobler, Mathias Wi—22(1):615 
Townsend, John F—22(2):873 
Triplett, J.K—22(1):79 
Tumlison, Renn—22(1):749 
Turner, Billie L—22(2):1 249 


Vandervoort, Carla—22(1):265 
Vazquez, M._Lucfa—22(2):1091 
Ward, Daniel B.—22(2):1111 
Weakley, A.'S.—22(1):79 

Werier, David A.—22(2):1049 
Whittemore, Alan T.—22(1):769 


1266 


Wilder, George J.—22(1):/87 
Witsell, C. Theo—22(1):801 
Wofford, B.Eugene—22(1):447 


BRIT.ORG/SIDA 22(2) 


Woolbright, Joe—22(1):801 
Wunderlin, Richard P—22(1):97, 635 
Wurdack, Kenneth J—22(1):469 


Zuloaga, Fernando O.—22(2):895 


BOTANICAL NAMES AND SuBJECT INDEX: VOLUME 22 (2006) 
New names (97) in bold face 


Achnatherum arnowiae—22(1):496 
Acrocystis—22(2):1049 
Ageratina thyrsiflora—22(2):1250 
Alternaria (fungus) —22(1):755 
Amaranthus blitum (Arkansas record)— 
22(1):817 
Amblysperma spathulata—22(1):539 
Amelanchier 
canadensis—22(2):1027 
obovalis—22(2):1027 
Amelichloa—22(1):146 
ambigua—22(1):147 
brachychaeta—22(1):147 
brevipes—22(1):147 
caudata—22(1):148 
clandestina—22(1):148 
hi Wi = 


—22(1):496 
Antennaria—22(1):561 
Apiaceae—22(2):955 
Apocynaceae—22(1):333, 355, 533:22(2):941 
Apocynoideae—22(1):333, 355 
Appalachian Mountains—22(1):79, 873 
Arizona—22(1):661 
ArkansaS—22(1):725, 817, 749; 22(2):801 
Arundinaria—22(1):79 
appalachiana—22(1):88 
gigantea—22(1):79 
falcata var. glomerata—22(2):1083 
Asclepiadoideae—22(1):533, 941 
Asphondylia borrichiae (midge) —22(1):755 
Aspidosperma 
rigidum—22(1):363 
schultesii—22(1):364 
Aster amellus—22(2):1087 
Asteraceae —22(1):367, 503, 539,545, 735, 755, 
22(2):866, 1075, 1087, 1249 
Aveninae—22(2):895 


SIDA 22(2): 1266. 2006 


Bahia, Brasil—22(1):333, 927 
Bahiella—22(1):342 
blanchetii—22(1):342 
infundibuliflora—22(1):345 
Bamboos—22(1):331 
Bambusoideae—22(1):79, 331, 1081 
Bauhinia —22(1):97 
amatlana—22(1):99 
andrieuxiim22(1):101 
arborea—22(1):102 
augusti—22(1):104 
ayabacensis—22(1):105 
beguinotii—22(1):107 
var. gorgonae—22(1):108 
brachycalyx—22(1):107 
Cookii—22(1):110 
geniculata—22(1):111 
haughtii—22(1):113 
petiolata—22(1):114 
var. caudigera—22(1):115 
picta—22(1):116 
section Amaria—22(1):97 
seleriana—22(1):117 
seminarioi—22(1):118 
Series Stenanthae—22(1):1 20 
stenantha—22(1):121 
weberbaueri—22(1):120 
Big Sandy Creek Unit (Big Thicket, Texas) — 
22(1):705 
Big Ticket National Preserve (Texas) —22(1):705, 
22(2):1175 
Big-Fruited Buckthorn—22(1):243 
Black Sapotes—22(1):277 
Boltonia 
asteroids var. asteroides—22(2):884 
caroliniana—22(2):884 
montana—22(2):874 


INDEX 


Borinda 
angustissima—22(1):332 
contracta—22(1):332 
nujiangensis—22(1):332 
utilis —22(1):332 

Borrichia frutescens—22(1):755 

Bradburia—22(1):503 

Brazil—22(1):123 (Sao Paulo), 333 

Bromeae—22(1):555,915 

Bromus—22(1):555 
ayacuchensis—22(2):919, 922 
berteroanus—22(2):922 
catharticus—22(2):923 

var. elata—22(1):556 
cebadilla—22(2):922 
coloratus—22(2):923 
flexuosus—22(2):923 
lanatus—22(2):923 
modestus—22(2):923 
pitensis—22(2):923 
segetum—22(2):923 
Striatus—22(2):922 
villosissimus—22(2):923 

Bryophyta—22(1):547, 549, 959 


Cactaceae—22(1):565 
Cacteae—22(1):565 
Caddo Parish (Louisiana) —22(2):1193 
Caesalpinoideae—22(1):97 
California—22(2):85 7 
Calyptranthes manuensis—22(2):93 1 
Cameron County (Texas) —22(2):1239 
Capraria mexicana—22(2):1 239 
Carex 

bicknellii—22(1):801 


gigantea X Carex lupuliformis—22(1):787 


reznicekii—22(2):1050 

stipata—22(2):1 221 
Carolinas—22(1):813 
Caryophyllaceae—22(2):1 145 
Cayratia japonica—22(1):813 
Central AmMerica—22(1):277 
Ceratochloa—22(1):555 
Cercideae—22(1):97 
Chamaesyce 

geyeri—22(2):1 221 


SIDA 22(2): 1267. 2006 


1267 


hypercifolia (Arkansas record) —22(1):817 
opthalmica (Arkansas record) —22(1):817 
Chaptalia hintonii—22(1):539 
Chelonanthus viridiflorus—22(1):137 
Chromolaena bigelovii—22(2):1 252 
Chromosome numbers—22(1):561 
Chrysopsidinae—22(1):503 
Chrysopsis—22(1):503 
Cicendia quandrangularis—22(1):1 33 
Clematis—22(1):65 
addisonii—22(1):65 
carrizoensis—22(1):67 
glaucophylla—22(1):65 
morefieldiim22(1):821 
texensis—22(1):65 
versicolor—22(1):65 
Collier County (Florida) —22(1):787 
Convolvulaceae—22(1):151,177, 197, 209 
Costa Rica—22(2):1071 
Crataegus 
ashei—22(2):1012 
austromontana—22(2):1016 
brittonii—22(1):423 
eximia—22(2):973 
gilva—22(2):973 
harbisonii—22(2):1012 
mendosa—22(2):973 
opima—22(2):973 
pallens—22(2):973 
pinetorum—22(2):973 
pulcherrima—22(2):973 
sargentii—22(2):973 
series Bracteatae—22(2):1009 
series Parvifoliae—22(1):423 
series Pulcherrimae—22(2):973 
series Triflorae—22(2):1009 
tecta—22(2):973 
triflora—22(2):1018 
uniflora—22(1):423 
X vailiae—22(1):423 
venusta—22(2):973 
Croptilon—22(1):503 
Croton alabamensis—22(1):469 
Cuban Herbaria—22(1):533 
Curtia tenuifolia—22(1):133 
Cuscuta—22(1):151 


1268 


californica—22(1):1 77 
campestris—22(1):15] 
coryli—22(1):209 
glabrior—22(1):151 
gronovii—22(1):197 
harperi—22(1):151 
howelliana—22(1):177 
indecora—22(1):209 
var. attenuata—22(1):216 
jepsonli—22(1):209 
occidentalis —22(1):177 
pentagona—22(1):151 
plattensis—22(1):151 
polygonorum—22(1):151 
runyonii—22(1):151] 
Salina—22(1):177 
sandwichiana—22(1):151 
Subinclusa—22(1):1 77 
Suksdorfii—22(1):1 77 
umbrosa—22(1):197 
warneri—22(1):209 
Cymopterus 
breviradiatus—22(2):955 
davidsonii—22(2):956 
glomeratus var. fendleri—22(2):956 
sessiliflorus—22(2):956 
Cyperaceae—22(1):/81,801,22(2): 1049 
Cyperus 
nigromarginata—22(2):1049 
umbellata—22(2):1049 
Dahlia—22(1):545 
sublignosa—22(1):545 
Delaware—22(1):825; 22(2):1027 
Delmarva Peninsula—22(1):825;22(2):1027 
Dendrophorbium 
paranense—22(1):127 
restingae—22(1):123 
Diospyros 
conzattii—22(1):287 
costaricensis—22(1):291 
gomeziorum—22(1):282 
pergamentacea—22(1):297 
riojae—22(1):279 
tuxtlensis—22(1):295 


SIDA 22(2): 1268. 2006 


BRIT.ORG/SIDA 22(2) 


Dodecatheon—22(1):461; 22(2):863 
austrofrigidum—22(1):462 
clevelandii 

var. gracile—22(2):863 

var. insulare—22(2):863 

var. patulum—22(2):863 
frigidum—22(1):461 
pulchellum 

var, Macrocarpum—22(2):863 

var. Shoshonense—22(2):864 
utahense—22(2):864 

Drepanostachyum 
falcatum—22(2):1084 

var. sengteeanum—22(1):332; — 
22(2):1081 


Ebenaceae—22(1):277 
Ecuador—22(2):931 
Eichhornia crassipes—22(1):749 
Eleocharis liesneri—22(1):781 
x Elyleymus 
hultenii—22(1):496 
mossii—22(1):497 
ontariensis—22(1):497 
Elymus—22(1):485 
xX cayouetteorum—22(1):498 
churchii—22(1):486 
hystrix—22(1):489 
lanceolatus subsp. riparius—22(1):498 
pringlei—22(1):485 
texensis—22(1):488 
Endophytes—22(1):755 
Ericameria nauseosa subsp. ammophila— 
22(2):868 
Erigeron strigosus—22(1):265 
var. traversii—22(1):273 
Eriogonum 
callistum—22(2):857 
Sect. Lanocephala—22(2):857 
Erodium malacoides—22(2):1 245 
Espiritu Santo Island—22(2):1135 
Eugenia 
haberi—22(2):1071 
yasuniana—22(2):934 
Eupatorieae—22(2):1249 
Eupatorium madrense—22(2):1253 


INDEX 


Euphorbia graminea (Arkansas record)— 
22(1):817 

Euphorbiaceae—22(1):469 

Eurytaenia texana—22(2):1221 

Exotic Plant Introductions (Kansas) —22(1):777 


Fabaceae —22(1):97 

Fargesia apicirubens—22(1):331 

Feral Goats—22(2):1135 

Ficus 
palmata—22(1):/769 
pseudos-carica—22(1):769 

Florida—22(1):635, 787 
Botanist—22(1):607 

Fraser, John (Folio) —22(2):1111 


Galium tricornutum (Oklahoma record)— 
22(2):1235 
Gentiana prostrata—22(1):133 
Gentianaceae—22(1):129 
Gentianella 
cosmantha—22(1):135 
fabrisii—22(1):130 
magellanica—22(1):135 
multicaulis—22(1):136 
ottonis—22(1):136 
Geocarpon minimum—22(2):1145 
Georgia—22(1):227, 243 
Geraniaceae—22(2):1245 
Gerbera—22(1):539 
Goldenasters—22(1):503 
Gonolobinae—22(1):533 
Gratiola brevifolia—22(1):825 
Green Fringed Orchid—22(1):805 
Gulf of California, Mexico—22(2):1135 


_ 


Hamamelidaceae—22(2):849 

Hamamelis—22(2):849 
macrophylla—22(2):855 
ovalis—22(2):850, 854 
vernalis—22(2):854 
virginiana—22(2):854 

Hardin County (Texas) —22(2):1175 

Hedyotis 
australis—22(1):831 
corymbosa (Arkansas record) —22(1):81 7 


SIDA 22(2): 1269. 2006 


1269 


Heterotheca—22(1):503 
depressa—22(1):379 
horrida—22(1):378 
polothrix—22(1):373 
polothrix—22(1):379 
sierrablancensis—22(1):374, 379 
stenophylla—22(1):377 
villosa—22(1):367 

var. angustifolia—22(1):376 

Hexalectris grandiflora—22(2):1 239 

Hill Cane—22(1):79 

Hispaniola—22(2):941 

Jatropha cinerea—22(2):1142 

India—22(2):935 

Indocalamus hamadae—22(1):332 

Insect Galls—22(1):/55 

Ipomoea quamoclit (Arkansas record)— 

22(1):81/7 

Kansas—22(1):777 

Koanophyllon 
palmeri—22(2):1251, 1252 
solidaginifolium—22(2):1252 


Lance Rosier Unit (Big Thicket, Texas)— 
22(2):1175 

Lantana—22(1):381 

camara—22(1):381 

subsp. aculeata—22(1):394 

nivea subsp. mutabilis—22(1):395 

strigocamara—22(1):392 
Lee County (Florida) —22(1):787 
Leibnitzia—22(1):539 
Lespedeza thunbergii—22(1):/77 
xLeydeum littorale—22(1):498 
Leymus californicus—22(1):498 
Lilaeopsis schaffneriana ssp. recurva—22(1):661] 
Lobatae (Quercus) —22(2):1091 
Lophophora williamsii—22(1):565 
Louisiana—22(1):805; 22(2):1193, 122] 
Madagascar—22(2):887 
Madre de Dios, Peru—22(1):615 
Mahler, Wm. F—22(2):847 
Maloideae—22(2):1027 
Mandevilla 

angustata—22(1):356 


1270 


boliviensis—22(1):359 
tristis—22(1):359 
vanheurckii—22(1):362 

Maryland—22(2):1027 

Matelea—22(2):941 
corynephora—22(2):942 
phainops—22(2):948 
rhynchocephala—22(2):949 
torulosa—22(2):951 

McFarlin, James Brigham—22(1):607 

Medicinal Plants —22(1):593 

Melanthiaceae—22(1):447 

Mexican Red Oaks—22(2):1091 

Mexico—22(1):277; 22(2):904, 1119 (central) 

Mississippi—22(2):849 

Missouri (record) —22(1):831; 22(2):1145 

(southwest) 

Monteverde Region (Costa Rica) —22(2):1071 

Moraceae—22(1):769 

Morus nigra—22(1):/69 

Mutisieae—22(1):539 

Myagrum perfoliatum—22(1):777 

Myrtaceae —22(2):927,931, 1071 


New Mexico (records) —22(2):1225 
North America Grasses—22(1):495 
North Carolina—22(1):813 
North-South Transition of Flora—22(1):725 
Noticastrum—22(1):503 
Nymphaea minuta—22(2):887 
Nymphaeaceae—22(2):887 
Odontadenia killipii—22(1):362 
Ohio—22(1):765 
Oldenlandia—22(1):305 
boscii—22(1):305 
corymbosa—22(1):305 
drymarioides—22(1):305 
lancifolia—22(1):305 
microtheca—22(1):305 
ovata—22(1):305 
pringlei—22(1):305 
salzmannii—22(1):305 
uniflora—22(1):305 
Opuntia tapona—22(2):1139 
Orchidaceae—22(1):805, 935 


SIDA 22(2): 1270. 2006 


BRIT.ORG/SIDA 22(2) 


Oregon—22(1):461 
Osbertia—22(1):503 


Pakistan —22(1):593 
Palm swamp forests—22(1):615 
Palma Real River—22(1):615 
Pantling, Robert—22(2):935, 939 (portrait) 
Parentucellia viscosa (Oklahoma record)— 
2212 )51 35 
Parietaria floridana 
22(1):817 
Pasco County, Florida—22(1):635 
x Pascoleymus—22(1):499 
bowdenii —22(1):499 
Pediomelum—22(1):227 
esculentum—22(2):1222 
piedmontanum—22(1):229 
Pennellia robinsonii—22(1):661 
Peru—22(1):615;22(2):915, 931 
Peyritschia 
conferta—22(2):902 
deyeuxioides—22(2):902 
howellii—22(2):897; 902 
humilis—22(2):902 
koelerioides—22(2):901 
pinetorum—22(2):899; 901 
pringlei—22(2):902 
Phacelia sonoitensis—22(1):661 
Piedmont Physiographic Province —22(1):227 
Plantaginaceae—22(1):825 
Platanthera lacera—22(1):805 
Poa 
annua—22(2):912 
bigelovii—22(2):912 
compressa—22(2):913 
fendleriana ssp. albescens—22(2):913 
ssp. fendleriana—22(2):913 
infirma—22(2):912 
matri-occidentalis—22(2):906 
subsp. mohinorensis—22(2):911 
mulleri—22(2):91 3 
orizabensis—22(2):91 2 
pratensis—22(2):913 
strictiramea—22(2):913 
Poaceae—22(1):79, 145, 485, 495, 555: 
22(2):1081, 895, 904,915 


(Arkansas record)— 


INDEX 


Poeae—22(2):904 
Poinae—22(2):904 
Polygonaceae—22(2):857 
Polytrichaceae—22(1):547 
Polytrichastrum alpinum var. sylvaticum— 
22(1):547 

Pontederiaceae—22(1):749 
Pooideae—22(2):895,915 
Prairie Fleabane—22(1):265 
Primulaceae—22(1):461, 863 
Pseudoroegneria spicata 

f.inermis—22(1):499 

f. pubescens—22(1):499 
Psidium cauliflorum—22(2):927 
Punjab, Pakistan —22(1):593 


Quercus —22(2):1091 


Rabbitbrush—22(2):866 

Ranunculaceae —22(1):65, 821 

Rhynchosia minima—22(1):661 

Ridge and Valley Physiographic Province— 
22(2):873 

Rosaceae—22(2):973, 1009, 1027 

Rubiaceae —22(1):831 


San Luis Valley (Colorado) —22(2):866 

San Rafael State Park (Arizona) —22(1):661 

Sapotes, Black—22(1):277 

Schoenoplectus hallii—22(2):1159 

Schultesia guianensis—22(1):140 

Scrophulariaceae—22(1):3811 

Senecioneae—22(1):1 23 

Seymeria falcata—22(1):811 

Shinners, Lloyd H—22(2):847 

Short’s Goldenrod—22(1):735 

Sibara virginica—22(1):661 

Sida, Contributions to Botany —22(2):847 

Sideroxylon—22(1):243 
lanuginosa—22(1):243 
macrocarpum—22(1):245 

Solidago shortii—22(1):735 

Sonoita Creek State Natural Area (Arizona)— 

22(1):661 

South America—22(1):781 

South Carolina—22(1):227 

Southwestern Arkansas—22(1):749 


SIDA 22(2): 1271. 2006 


1271 


Southwestern U.S.A—22(1):769, 955 
Sphagnaceae—22(2):959 
Sphagnum 
beothuk—22(2):966 
bergianum—22(2):964 
kenaiense—22(2):961 
mcqueenii—22(2):959 
sitchense—22(2):969 
talbotianum—22(2):970 
Starkey Wilderness Preserve—22(1):635 
Stenanthium diffusum—22(1):450 
Stevia salicifolla—22(2):1 249 
Stipeae—22(1):145 
Symphyotrichum patens—22(2):1075 
var. gracile—22(2):1076 
var, patens—22(2):1076 
var, patentissinum—22(2):1075 


Talinum calycinum—22(2):1 222 
Tehachapi Mountains—22(2):857 
Tennessee—22(1):447, 821 
Tetraphidaceae—22(1):549 
Tetraphis pellucida—22(1):551 
var. trachypoda—22(1):551 
Texas—22(1):65 (northeastern), 265, 705, 811; 
22(2)31 175, 1239, 1259, 1245 
Thieret, John W—22(1):1,3, 21,25, 33,51,55 
Tomentaurum—22(1):503 
Trichocline—22(1):539 
hieracioides—22(1):539 
Trichome morphology—22(2):1091 
Type Localities of Ohio Plants—22(1):765 


United Status (record) —22(1):811 


Venezuela—22(1):355 
Verbenaceae—22(1):381 
Viola 

grahamii—22(2):1119 

hookeriana—22(2):1119 
Violaceae—22(2):1119 
Viorna (subgenus)—22(1):65 
Virginia (Eastern) —22(2):1027 
Vitaceae—22(1):813 


Walter, Thomas—22(2):1111 
Washington—22(1):461 
Water Hyacinth—22(1):749 


— 


1272 
Waterlily—22(2):887 


West Indies —22(1):533 
Witch-Hazel—22(2):849 


BRIT.ORG/SIDA 22(2) 


Zeuxine 
goodyeroides—22(2):935 
pantlingii—22(2):935 

Zygostigma australe—22(1):1 36 


97 New NAMES AND NEw COMBINATIONS: 
Votume 22 (2006) 


Achnatherum arnowiae (S.L. Welsh & N.D. At- 
wood) Barkworth, comb. nov.—22(1):496 
Amelichloa Arriaga & Barkworth, gen. nov.— 
22(1):146 

Amelichloa ambigua (Speg.) Arriaga & 
Barkworth, comb. nov.—22(1):147 

Amelichloa brachychaeta (Godr.) Arriaga & 
Barkworth, comb. nov.—22(1):147 

Amelichloa brevipes (£. Desv.) Arriaga & 
Barkworth, comb. nov.—22(1):147 

Amelichloa caudata (Trin.) Arriaga & Barkworth, 
comb. nov.—22(1):148 

Amelichloa clandestina (Hack.) Arriaga & 
Barkworth, comb. nov.—22(1):148 

Ammophila breviligulata subsp. 
champlainensis (F. Seym.) Walker, Paris & 
Barrington ex Barkworth, comb. nov.— 
22(1):496 

Arundinaria appalachiana Triplett, Weakley & 
L.G. Clark, sp. Nov.—22(1):88 


Bahiella J.F Morales, gen. nov.—22(1):342 

Bahiella blanchetii (A. DC.) J.- Morales, comb. 
nov.—22(1):342 

Bahiella infundibuliflo 
22(1):345 

Bauhinia amatlana Wunderlin, sp. nov.— 
22(1):99 

Bauhinia arborea Wunderlin, sp. nov.— 
22(1):102 

Bauhinia ayabacensis Wunderlin, sp. nov.— 
22(1):105 

Bauhinia geniculata Wunderlin, sp. nov.— 
22(1):1 11 

Bauhinia petiolata var. caudigera (S.F Blake) 
Wunderlin, comb. nov.—22(1):115 

Boltonia montana JF Townsend & V. Karaman- 


Castro, Sp. NOV.—22(2):874 


).F. Morales, sp.nov,.— 


SIDA 22(2): 1272. 2006 


Borinda angustissima (1.P.Yi) Stapleton, comb. 
NOV.—22(1):332 

Borinda contracta (T.P. Yi) Stapleton, comb. 
NOV.—22(1):332 

Borinda nujiangensis (Hsueh & C.M. Hui) 
Stapleton, comb. nov.—22(1):332 

Borinda utilis (TP. Yi) Stapleton, comb. nov.— 
22(1):332 

Bromus ayacuchensis Saarela & P.M. Peterson, 
Sp.nov.—22(2):919 

Bromus catharticus var. elata (£. Desv.) 

juelo, comb. nov.—22(1):556 


— 


Plane 


Calyptranthes manuensis 8. Holst & ML. 
Kawasaki, Sp NOv.—22(2):93 | 

Carex reznicekii Werier, 5p. nov.—22(2):1050 

Clematis carrizoensis D. Estes, sp. nov.— 
22(1):67 

Cuscuta indecora var. attenuata (Waterfall) 
Costea, comb. & stat. nov.—2.2(1):216 

Cymopterus breviradiatus (W.L. Theob. & C.C. 
Tseng) R.L. Hartm., comb. & stat. nov.— 
22(2):955 

Cymopterus davidsonii (J.M.Coult.& Rose 
Hartm., comb. nov.—22(2):956 

Cymopterus glomeratus var.fendleri (A. Gray) 
R.L. Hartm., comb. & stat. nov.—22(2):956 

Cymopterus sessiliflorus (W.L. Theob. & C.C. 
Tseng) R.L. Hartm., comb. nov.—22(2):956 


Dahlia sublignosa (P.D.Sarensen) D.E.Saar & PD. 
S@rensen, comb. & stat. nov.—22(1):545 

Dendrophorbium restingae A Jeles, J. N.Nakaj. 
& Stehmann, sp. nov.—22(1):1 23 

Diospyros costaricensis M.C. Provance & A.C. 
Sanders, sp. nov.—22(1):291 

Diospyros gomeziorum M.C. Provance & AC. 
Sanders, sp. nov.—22(1):282 


— 


R.L. 


INDEX 


Diospyros tuxtlensis M.C. Provance & A.C. Sand- 
ers, SO. NOV.—22(1):295 

Dodecatheon austrofrigidum K.L. Chambers, 
sp. NOV.—22(1):462 

Dodecatheon clevelandii var. gracile (Greene) 
Reveal, comb. nov.—22(2):863 

Dodecatheon clevelandii var. insulare (HJ. 
Thomps.) Reveal, comb. nov.—22(2):863 

Dodecatheon clevelandii var. patulum 
(Kuntze) Reveal, comb. nov.—22(2):863 

Dodecatheon pulchellum var. macrocarpum 
(A. Gray) Reveal, comb. nov.—22(2):863 

Dodecatheon pulchellum var. shoshonense 
(A. Nelson) Reveal, comb. nov.—22(2):864 

Dodecatheon utahense (N.H. Holmgren) Re- 
veal, stat. Nov.—2.2(2):864 

Drepanostachyum falcatum 
sengteeanum Stapleton, var. nov.— 
22(1):332 


Val. 


xX Elyleymus hultenii (Melderis ex Hulten) 
Barkworth, comb. nov.—22(1):496 

Elyleymus mossii (Lepage) Barkworth, comb 
nov.—22(1):497 

Elyleymus ontariensis (Lepage) Barkworth, 
comb. nov.—22(1):497 

Elymus Xcayouetteorum (Boivin) Barkworth, 
comb. nov.—22(1):498 

Elymus churchii J.J.N. Camp., sp. nov.— 
22(1):486 

Elymus lanceolatus subsp. riparius (Scribn. & 
J.G. Smith) Barkworth, comb. & stat nov.— 
22(1):498 

Elymus texensis J.J.N. Camp., sp. nov.— 
22(1):488 

Ericameria nauseosa subsp. ammophila L.C. 
Anderson, subsp. nov.—22(2):868 

Erigeron strigosus var. traversii (Shinners 
Noyes, comb. & stat nov.—22(1):273 

Eriogonum callistum Reveal, sp. nov.— 
22(2):857 

Eriogonum Sect. Lanocephala Reveal, sect. 
Nov.—22(2):857 

Eugenia haberi Barrie, sp. nov.—22(2):1071 


pale’, 


SIDA 22(2): 1273. 2006 


1273 


Eugenia yasuniana B. Holst & M.L. Kawasaki, sp. 
nov.—22(2):934 


Fargesia apicirubens Stapleton, sp. nov.— 
22(1):331 

Gentianella fabrisii Filippa et Barboza, sp. 
nov.—22(1):130 

Hamamelis ovalis S.W. Leonard, sp. nov.— 
22(2):850 

Heterotheca polothrix Nesom, nom. et stat. 
nov.—22(1):373 

Heterotheca sierrablancensis (Semple) 
Nesom, comb. & stat. nov.—22(1):374 


Indocalamus hamadae (Hatus.) Stapleton, 
comb.nov.—22(1):332 


Lantana camara subsp. aculeata (L.) R.\W.Sand- 
ers, comb. & stat. nNOV.—22(1):394 

Lantana nivea subsp. mutabilis (WJ. Hook.) 
RW. Sanders, comb. & nov.—22(1):395 

Lantana strigocamara R.W. Sanders, sp. nov.— 
22(1):392 

xX Leydeum littorale (H.J. Hodgs.& W.W. Mitch.) 
Barkworth, comb. nov.—22(1):498 

Leymus californicus (Bol. ex Thurber) 
Barkworth, comb. nov.—22(1):498 


Mandevilla angustata (Steyerm.) J.F. Morales, 
comb. & stat. nov. —22(1):356 

Mandevilla tristis J.F. Morales, sp. nov.— 
22(1):359 

Matelea corynephora Krings, sp. nov.— 
22(2):942 

Matelea phainops Krings, sp. nov—22(2):948 

Matelea rhynchocephala Krings, sp. nov.— 
22(2):949 

Matelea torulosa Krings, sp. nov.—22(2):951 

Nymphaea minuta K. Landon, R.A. Edwards & 
P|. Nozaic, sp. nOV.—22(2):887 


x Pascoleymus Barkworth, gen. hybr. nov.— 
22(1):499 

x Pascoleymus bowdenii (Boivin) Barkworth, 
comb. nov.—22(1):499 

Pediomelum piedmontanum JR. Allison, M.W. 
Morris & A.N. Egan, sp. nov.—22(1):229 


1274 


Peyritschia howellii (Hitchc.) Finot & PM.Peter- 
son, comb. nov.—2.2(2):897 

Peyritschia pinetorum (Swallen) Finot & P.M. 
Peterson, comb, nov.—22(2):899 

Poa matri-occidentalis P.M.Peterson & Soreng, 
Sp. NOV.—22(2):906 

Poa matri-occidentalis subsp. mohinorensis 
soreng & PM. Peterson, subsp. nov.— 
22(2):911 

Polytrichastrum alpinum var. sylvaticum 
(Menzies) G.L. Merrill, comb. nov.—22(1):547 

Pseudoroegneria spicata f. inermis (Scribn. & 
J.G. Sm.) Barkworth, comb. nov. & stat. nov.— 
22(1):499 


~ 


AL ta f kK (Cl 


Barkworth, comb. nov. & stat.nov.—22(1):499 
Psidium cauliflorum Landrum & Sobral, sp. 
nov.—22(2):927 
Psidium cauliflorum Landrum & Sobral, sp. 
NOV.—22(2):931 


SIDA 22(2): 1274. 2006 


BRIT.ORG/SIDA 22(2) 


Sideroxylon macrocarpum (Nutt.) J.B. Allison, 
comb. nov.—22(1):245 

Sphagnum beothuk Andrus, sp. nov.— 
22(2):966 

Sphagnum bergianum Andrus, sp. nov.— 
22(2):964 

Sphagnum kenaiense Andrus, sp. nov.— 
22(2):961 

Sphagnum mcqueenii Andrus, sp. nov.— 
22(2):959 

Sphagnum sitchense Andrus, sp. nov.— 
22(2):969 

Sphagnum talbotianum Andrus, sp. nov.— 
22(2):970 

Stenanthium diffusum Wofford, sp. nov.— 
22(1):450 


Tetraphis pellucida var trachypoda (Kindb. ex 
Paris) Harpel, comb. nov.—22(1):551 

Zeuxine pantlingii Av. Bhattacharjee & HJ. 
Chowdhery, sp. nov.—22(2):935 


i A 


olume | of the Illustrated Flora of East Texas is the first fully 
Ne flora for East Texas, a species-rich area that 
contains roughly two-thirds of all the plant species known for 
Texas. The book covers all the native and naturalized ferns and 
similar plants, gymnosperms, and monocotyledons (1,060 
species) known to occur in East Texas. 

Published ape) Botanical Research Institute of Texas 

clo WMusriated ts plofas ro oje 


Ilak . €7; RIT 
Texas (BRIT 


and the 
Austin College Center for Environmental Studies. 
www.easttexasflora.org 


2 vy" x10 ve 


1,614 pages 
1,200+ bist Soules 


1,000+4 


Sida, Botanical Miscellany 126 Illustrated Flora 


issn 0833-1475 of East Texas 
isbn 1-889878-12-X, hbk. 
George Diggs, Barney Lipscomb, 
May 2006 Monique Reed, Robert J. O’Kennon 
Big “D" to the Big Thicket there is a land 
ee interwoven with groves of timber, 
stately pines, hardwoods, wed) ROnEYsuCNG a 
hid: d, d: 


€20905 


please inquire) 


Available from Botanical Research Institute of Texas Press 


509 Pecan Street 
md VAL, LL 761 4060, U.S.A | 


E-mail: dies 
Fax: 1-81 Bar aie aoe 


Pee nC ie sear 1 EA 9 


BRIT 


TS photographic guide to the Wild Flowers of 
WILD FLOWERS OF MOMBACHO 
Mombacho is color-coded and arranged by Family NICARAGUA 
within the color sections. It covers flowering plants found 
on the whole of the Volcano; which includes both tropical 
cloud forest and dry, deciduous forest, at lower altitudes. 
The book covers 200 plants, each of which has two or 
three photographs and a short description in both English 
and Spanish giving common name(s); a short description 
of the plant; height; flowering period; habitat and any 
known use of the plant. The vast majority of plants are 


likely to be found in similar habitats throughout tropical Ss HELEN PACKERING 


America and nearly one third are pan tropical, making this 


book of interest to a wide audience throughout the tropics. WILD FLOWERS OF MOMBACHO 
ICARAGUA 
Sida, Bot. Misc. No 28, 2006 lca 
issn 0823-1475 FLORES SILVESTRES DEL MOMBACHO 
isbn 1-889878-14-6 BY HELEN PICKERING 
6.5" x 8.5", x + 217pp. 
over 500 photographs ) 
5 + p&p" 
SA $3 


“International: $7 BI 
Texas residents add $1.24 sales tax PRESS 


UISIANA TEXAS. GEORGIA NORTH. 


\N AHOMA. VIRGINIA OREGON. NEBRASKA ALABAMA NEW MEXICO ea 


ITZERLAND “ze 


YA JORDON ‘en : 


XACRUZ ME NOY 


ALGO GUADALAJARA od COAHUILA YUCATAN Sih